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The Gardens 1 Bulletin 

Singapore 




NATIONAL PARKS BOARD 

Singapore Botanic Gardens Cluny Road Singapore 259569 Tel: 4741165 Telefax: 4754295 



THE GARDENS' BULLETIN 



The Gardens' Bulletin Singapore publishes papers on plant taxonomy (including revisions), 
horticulture, phytogeography, floristics, morphology, anatomy and related fields with 
emphasis on plants in the West Malesian region. 



EDITORIAL BOARD 



Dr Ruth Kiew 
(Editor) 

Singapore Botanic Gardens 

Dr T.W. Foong 
(Assist. Editor) 
Singapore Botanic Gardens 

Dr S.C. Chin 

Singapore Botanic Gardens 

Dr M.J.E. Coode 
Royal Botanic Gardens 
Kew, U.K. 



Dr R.T. Corlett 
University of Hong Kong 
Hong Kong 

Dr M.C. Roos 
Rijksherbarium 
Leiden, Netherlands 

Dr E. Soepadmo 

Forest Research Institute Malaysia 
Kepong, Malaysia 

Dr W.K. Tan 

Singapore Botanic Gardens 



The Gardens' Bulletin is published twice yearly by the National Parks Board, Singapore. 
Neither the National Parks Board nor the Editorial Board is responsible for the opinions or 
conclusions expressed by the contributing authors. 

The annual subscription for the Gardens' Bulletin is Singapore $100.00 including 
postage. Overseas subscribers are required to make payment in the form of bank drafts or 
international money orders in Singapore currency payable to National Parks Board, 
Singapore. 

Instructions for contributing authors are found on the inside backcover. 



M 

1 

The Gardens 1 Bulletin 

Singapore 



VOL. 53 (Parts 1 & 2) July 2001 ISSN 0374-7859 

CONTENTS 

Argent, G. 

Contributions to the flora of Mount Java VI. A new banana species. Musa johnsii 
(Musaceae) from New Guinea 1 

Bastmeijer, J.D. and R. Kiew 

A new Cryptocoryne hybrid (Araceae) from the Bukit Timah Nature 

Reserve. Singapore 9 

Boyce, P.C. 

The genus Rhapiduphora Hassk. (Araceae-Monsteroideae- 

Monstereae) in Borneo 19 

Boyce, P.C. 

The genus Rhaphidophota Hassk. (Araceae-Monsteroideae- 
Monstereae) in New Guinea. Australia and the tropical Western 

Pacific 75 

Jong, K. and G. Argent 

Cytology of two new species of Musa (Musaceae) from Borneo and 

their sectional relationship 185 

Khaw, S.H. 

The genus Etlingera (Zingiberaceae) in Peninsular Malaysia 

including a new species 191 

Kiew, R. 

The limestone begonias of Sabah. Borneo - flagship species for 

conservation 241 

Kiew, R. 

Begonia conipila Irmsch. ex Kiew (Begoniaceae) from the Gunung 

Mulu National Park. Sarawak. Malaysia 287 

Ng, F.S.P. 

New species, varieties and reductions in Diospyros (Ebenaceae) in Borneo and 
Peninsular Malaysia including peninsular Thailand 291 




Tan, B.C. and E. H. Mandia 

New and noteworthy records of mosses from Mindoro, the 

Philippines, and their biogeographical implication 315 

Turner, I. M. 

New combinations of Malesian Myricaceae 323 

Wong, C, R. Kiew, A. Lamb, Ohn Set, S. K. Lee, L. H. Gan and Y. Y. Gan 

Sectional placement of three Bornean species of Musa (Musaceae) based on 
Amplified Fragment Length Polymorphism (AFLP) 327 

Yii P.C. and P.P.K. Chai 

New combinations, new names and new species of Madhuca 

(Sapotaceae) from Sabah and Sarawak. Borneo 343 

Corrigenda - I. Salma. The significance of pollen morphology in the 

taxonomy of the genus Durio (Bombacaceae) 357 

BOOK REVIEW 

Secondary Forest Trees of Kalimantan Indonesia. A Manual for 300 

Selected Species F.S.P. Ng - 359 

Obituary Dr Gunnar Seidenfaden J. J. Vermeulen - 363 



Date of Publication: 31 July 2001 

Published by 

National Parks Board 
Singapore Botanic Gardens 
Cluny Road 
Singapore 259569 



Printed by Oxford Graphic Printers Pte Ltd 



Gardens' Bulletin Singapore 53 (2001) 1-7. 



Contributions to the Flora of Mount Jaya VI. 
A New Banana, Musa johnsii (Musaceae) from 
New Guinea 

G. ARGENT 
Royal Botanic Garden Edinburgh. EH3 5LR. U.K. 

Abstract 

A new species of banana, Musa johnsii Argent, is described and illustrated from Papua 
(formerly Irian Jaya). Indonesian New Guinea, and its possible position in the genus is 
discussed. 

Introduction 

New Guinea is an important centre of diversity for the genus Musa. With 
the addition of this new species, there are ten wild species recorded most 
of which are endemic. The area is also arguably the richest place in the 
world for indigenous cultivars (Simmonds 1966), many of which are diploids 
and potentially important as a gene reservoir. This new banana has all the 
attributes of a wild species in that it has a non-parthenocarpic seedy fruit 
and regenerates naturally from seed distributed by animals, yet it has a 
sterile, non-seedy terminal portion and was cultivated in at least one village 
(Utekini) for its use as a vegetable. 

Terminology and description follow previous tradition as used by 
Simmonds (1962, 1966) and Argent (1976). 

Musa johnsii Argent sp. nov. 

Species insignis propfer infructescentia densum paene sphaericum fructuum 
subsessilium primo ad apices truncatis demum irregulariter schizocarpicis. 
Ab omnibus species Musae differt fructu maturo cum tertia parte distali 
medullae sterilis composita. 

St 

Typus: Indonesia: Papua. Freeport mining concession above Timika. 1 
Nov 2000 Argent et al. 00562 (holo BO; iso K, E). 

Fig 1 & 2. 

Clump forming herbaceous plant. Pseudostem to 4 m tall, 28 cm diam. 
near the base. Predominantly brown with dead clinging pseudostems. 



2 



Gard. Bull. Singapore 53 (2001) 




otherwise green with some dark brown coloration in the upper parts, no 
wax; undersheath cream to white, juice milky white or creamy. Suckering 
moderate, arising alongside or up to 20 cm from parent, erect or nearly so. 
Sucker leaves mostly auriculate. Shoulder brown or green, entire, smooth 
and appressed, without any scarious margin. Fourth last leaf 242 x 88 cm, 
right handed to c. 8 mm, (almost symmetrical), the base cordate to weakly 
auriculate. Other leaves often left-handed and predominantly strongly 
auriculate, all green, hardly different in colour above and below, slightly 
paler beneath and with the prominent midrib, mostly pale yellow, sometimes 
with a little brown proximally without obvious wax. Petiole 52 x 4.5 cm, 
the adaxial channel green, open with only slightly incurved or erect margins 
which are green, not or only narrowly (to c. 1 mm) scarious, abaxially the 
petiole dark chocolate brown TS ratio 0.78 (see Argent 1976). PB ratio 4 — 
5. 

Peduncle stout, green, glabrous, densely scarred. Bunch horizontal, diagonal 
or occasionally held completely vertically downwards. The female bracts 
lanceolate to 54 x 12 cm, yellow, shiny outside, dull yellow and slightly 
paler inside, acuminate the apical 12 cm with the margins strongly inrolled. 
completely and quickly deciduous. Female flowers with a few staminodes 



Musa johnsii 



3 




Figure 2. Musa johnsii. fertile features: A. habit of fruit bunch with male bud x V 4 . B.T.S. fruit 
x , C. L.S. fruit x V 3 , D. whole fruit x 2 / 3 , E. female bract after dehiscence x l / 2 , F. free tepal of 
female flower x 1. G. style and stigma of female flower x 1, H. compound tepal of female 
flower x 1,1. male flower x 1. J. free tepal of male flower x 1. K. stamen from male flower x 1, 
L. compound tepal of male flower x 1, M. style and stigma from male flower x 1. 



4 



Card. Bull. Singapore 53 (2001) 



mostly less than half the length of the style, free tepal flushed purple. 
Ovary trilocular, each locule with the ovules in two rows. 

Male peduncle growing vertically downwards. The male bud up to 14 x 11 
cm, shiny yellow, imbricate for c. 2 cm from the tip. Male bracts 13 x 7 cm, 
shiny rich yellow outside, slightly darker near the margins, the tips pale 
green, shiny yellow inside although becoming dull inside after falling, with 
broadly rounded, obtuse apices, lifting to a high angle c. 45° to the axis; 
after falling only recurved at the base not revolute from the apex or margins. 
Male flowers, two-rowed, falling in a group, cream, the free tepal translucent 
white, with a rounded, erose upper margin and no wrinkle, just over half 
as long as the compound tepal. Compound tepal cream with pale yellow 
apices, the two longest points with irregular papillose to subdenticulate 
margins. 

Fruit bunch dense, sub-spherical in shape to 17 x 18 cm. Fruit two-rowed, 
the second hand with 14 fruits. The fruits irregular, apparently ageotropic 
as they show no curvature in any part of the bunch, ripening orange but 
the surface becoming mostly blackish by full maturity, 8 — 9 x 2.5 — 3.5 cm, 
3-, 4- or 5-angled, broadly truncate at the apex with large scars up to 2 cm 
diam., splitting irregularly to reveal pale pinkish orange pith and similarly 
coloured or yellowish flesh around the seeds in the carpel chambers, the 
pith white at the base of the fruit; yellow latex exuding from the cut skin of 
the immature fruit. Pedicel c. 2 — 3 mm, the fruits almost sessile. Seeds 
dark brown, 4 — 5 mm diam., irregular with rounded angles and with a 
distinctly domed boss opposite the hilum which is c. 2 mm in diam. and 
vertically striate, also with a raised band on one side from hilum to boss. 

Vernacular name: Mel in the Amungkal language of the Amungme Tribe. 

Uses: The Amungme people eat this species raw as a vegetable, slicing the 
young stalks and mixing them with local Begonia leaves (nilmanep) and 
salt. 

Notes: Named in honour of Professor R.J. Johns who has made a lifetime 
study of the plants of New Guinea and who first drew my attention to this 

species. 

This new species was a very surprising find on the southern side of 
the main range north of Timika. It is very distinctive in the dense subspherical 
bunch of almost sessile fruits, which are rather irregularly schizocarpic, and 
are truncate at the apex. Unlike any other Musa species, it has a sterile 
mucilaginous pith chamber in the distal third of the mature fruit. 




Figure 3. Professor R.J. "Bob' Johns with the fruit bunch and male bud of Musa johnsii. 



6 



Card. Bull. Singapore 53 (2001) 



Two populations were seen. The first where it was semi-cultivated 
for the edible leaf stalks near Utekini below Tembagapura at about 1600 
m. The second was on the Darnell Ridge below the Hanekam Tunnel 
between 1000 — 1500 m altitude, where several groups of plants occurred 
along the roadside well away from any indigenous habitation. The largest 
of these populations of more than a hundred plants displayed all stages 
from abundant small seedlings to young clumps, plants flowering in the 
female phase and older ones with mature fruit and male inflorescences. 
They thus appear to be totally non-seasonal. 

The fruits are puzzling. The distal pith chamber could be an attractive 
lure to the fruit but, for a wild banana, the number of seeds in the very 
restricted loculi is very low (an average of 46 for the 5 fruits for which 
seeds were counted) compared with many hundreds in most other wild 
Musa species. The flesh of this species in either the pith chamber or 
around the seeds in the loculi did not seem to be particularly palatable and 
the local people are not reported to eat the fruit at any stage. Animals 
were certainly visiting the ripe fruits and removing the contents and many 
of the seeds were damaged in the process. The fact that the female bracts 
are completely deciduous is remarkable in that the mature fruits are tightly 
packed in a very dense bunch in the sense of Argent (1976). This is no 
doubt achieved because of the strong revolution of the bracts from both 
base and apex, which results in their falling in tight rolls. 

This species grows at the highest altitude of the three wild bananas 
encountered in the Freeport project area around Timika. Musa banksii F. 
Muell. was common in the lowland areas not too swampy and waterlogged. 
M. lolodensis Cheesman occurred mainly at the upper margin of the heath 
forest at about 500 m altitude, although one clump was seen growing with 
Musa banksii at less than 100 m. Both these species can be easily 
distinguished from M. johnsii even in the sterile state, as they both have 
broad scarious margins at the upper edge of the leaf sheath or 'shoulders' 
(Argent 1976). The intense green colour of the foliage and orange colour 
of the fruit indicate that this species probably belongs in section 
Australimusa. It would appear to be closely related to M. lolodensis. The 
seeds of the two species are remarkably similar. Apart from the vegetative 
difference mentioned, the fruit of these two species are very different. M. 
lolodensis has a lax bunch of distinctly long pedicellate fruit, which splits in 
a much more regular fashion on ripening than this new species. 

There was another wild banana reported from Mt Jaya, which could 
be Musa ingens Simmonds, but this was not seen by the author. It is very 
distinct from this new species as it has glaucous waxy pseudostems, a 
reflexed 'shoulder' and a long pedicillate fruit with much larger seeds 8 — 9 
mm in diameter. 



Musa johnsii 



Acknowledgements 

This work was made possible with the logistical and financial assistance 
from PT Freeport Indonesia and Rio Tinto in conjunction with Herbarium 
Bogoriense. Kebun Raya Bogor and The Biodiversity Centre. Cenderawasik 
University. Manokvvari. I am also grateful to the Royal Botanic Gardens. 
Kew for permission to join their expedition and to Prof R. J. Johns and Dr 
T. Utteridge for their companionship and tireless support in organising the 
expedition. Dr Carolyn Cook provided the local names and much valued 
help at Tembagapura. The Royal Botanic Garden. Edinburgh gave 
permission for me to participate in the fieldwork and Ms Christina Oliver 
ably interpreted the specimens, photographs and crude field drawings. I 
am indebted to Dr Robert Mill for the translation of the Latin diagnosis. 

References 

Argent G.C.G. 1976. The wild bananas of Papua New Guinea. Notes 
Royal Botanic Garden Edinburgh. 35: 77 — 114. 

Simmonds. N.W. 1962. The Evolution of the Bananas. Longmans, U.K. 

Simmonds. N.W. 1966. Bananas. 2 nd ed. Longmans, U.K. 



Gardens' Bulletin Singapore 53 (2001) 9-17. 



A New Cryptocoryne Hybrid (Araceae) from the Bukit 
Timah Nature Reserve, Singapore 

JAN D. BASTMEIJER 
Oude Roswinkelerweg 72. 7822 AG Emmen. The Netherlands 

AND 
RUTH KIEW 

Singapore Botanic Gardens, 1 Cluny Road. Singapore 259569 

Abstract 

A Cryptocoryne from the Bukit Timah Nature Reserve. Singapore, identified for many 
years as C. griffithii Schott. is recognized to be a new hybrid species: Cryptocoryne xtimahensis 
Bastmeijer. It is difficult to accurately determine its parents, but good candidates are C. 
nurii Furtado and C. cordata Griff., both native to Johore. Peninsular Malaysia. 

Introduction 

The first collections of Cryptocoryne from Singapore were made in 1890 
by H.N. Ridley (1904, 1907, 1925), the first director of the Botanic Gardens 
Singapore. Two species were recorded: C. griffithii Schott from several 
freshwater localities and C. ciliata (Roxb.) Schott from tidal areas. Today. 
C. griffithii is still present in the MacRitchie Reservoir and Nee Soon 
swamp forest but C. ciliata is reported to have disappeared from Singapore 
(Tan. 1995). The latter is, however, a widespread species, occurring from 
India to Papua New Guinea, while C. griffithii is a rather narrow endemic 
being known from the south of Peninsular Malaysia (Malacca and Johore), 
as well as in Singapore. However, today there may be only a few localities 
left in southern Johore of the Malaysian population. 

The main interest in Cryptocoryne is as an ornamental plant for 
aquascaping. This hobby of cultivating tropical plants and keeping fish in 
aquaria has become very popular in Europe, the USA, and Japan. From 
the 1950s. Singapore was the main port for shipping wild collected plants 
to these regions, and plant nurseries developed to satisfy this demand. 

The attention of the first author was drawn to the problem of the 
identity of the Bukit Timah plant by a photograph of C. griffithii on the 
website of the Singapore Science Centre from an article by Tan et al. 



10 



Card. Bull. Singapore 51 (2001) 



(1994). The photograph presented there of C. griffithii did not match the 
European idea of C. griffithii very well. The same photograph appeared in 
Foulis et al. (1998). In June 1999, there was an opportunity to study the 
plant in question more closely in the Bukit Timah Nature Reserve. We 
found the plant flowering abundantly and collected some specimens for 
further investigations. 

A Short History of Cryptocoryne griffithii Schott 

The identity of the real C. griffithii has been in dispute, especially because 
of the erroneously named drawing in Hooker (1900). Ridley (1904) pointed 
out that the drawing actually represented his newly described C. purpurea 
Ridl.. In 1920, Engler gave a good description of both species. De Wit 
(1961) pointed to this problem again, and he was fortunate to get live 
plants from Malacca, from where Griffith had originally collected his plants. 
A detailed description of the true C. griffithii together with a nice drawing 
by Ike Zewald is found in De Wit (1961, 1971, 1990) and which closely 
matches Schott's unpublished drawing in Vienna (W) of C. griffithii. 

One of the first commercial collectors of C. griffithii from Singapore 
was Y.W. Ong (pers. comm.), who collected the plants in the reservoirs 
and shipped them to Europe as far back as 1948. However, all publications 
in Europe on this plant in the hobby magazines we could find refer to C. 
purpurea. At that time there was also much confusion about C. cordata 
Griff., which was not well understood, see for example Hoedeman (1948), 
Wendt (1952), Oskam & Van Ramshorst (1954) and De Wit (1951, 1958a, 
b). The reason for the lack of notes on C. griffithii is probably very simple: 
the species is very difficult to grow and neither hobbyists nor professionals 
were able to grow them at that time. But from 1961 onwards, C. griffithii 
was at last a well-known species. 

Around the 1970s, taxonomy in Cryptocoryne seemed to be resolved 
(De Wit, 1971), but in the following years the opposite became true. 
Jacobsen (1982, 1987) suggested a hybrid nature for C. purpurea. Jacobsen 
& Bogner (1986, 1987) gave a detailed review on the Cryptocoryne of the 
Malay Peninsula. Today, with more collections available and many 
experiments on cultivation, it appears that within some groups there is a 
near continuous variation, making it in some extreme examples, more or 
less a matter of taste how to interpret a newly imported plant. Furthermore, 
polyploidy and natural hybrids have proved to be rather common in 
Cryptocoryne. Today we know a couple of C. griffithii-\\ke. plants that 
show some variation. A recent review on C. griffithii and Cryptocoryne in 
general is found in Bastmeijer (2000). 



Cryptocoryne xtimahensis from Singapore 



1 1 



With the variation of C. griffithii now understood, it is possible to be 
certain that the Bukit Timah plant is a distinct new hybrid differing, among 
other characters, from C. griffithii in its narrowly ovate leaves, the long 
apex of the spathe and large, irregular protuberances of the limb. The fact 
that its pollen fertility was less than 10% and that it had never been found 
in fruit strongly suggests that it is a hybrid. 

Cryptocoryne xtimahensis Bastmeijer notosp. nov. (Araceae) 

Lamina spathae protuberationibus irregularibus obtecta, longicaudata (in 
C. griffithii brevicaudata et dense verrucosa), basis laminae plus minusve 
auriculata (similiter in C. nurii), color laminae flavus protuberationibus 
atrorubis, fauce modice angusta (similiter in C. nurii). Folia eis speciei C. 
cordata simillima (sed C. cordata differt lamina spathae leavi tota flava) - 
Typus: Singapore, Bukit Timah Nature Reserve. Jungle Fall Stream. 18 
January 2001, dammed pools. Alt. c. 120 m., R. Kiew, S. Saifuddin, S. Teo 
& A.T. Gwee RK 5127. (holo SING; iso C. K). 

Rhizome creeping, OA — 0.8 cm diam.. stoloniferous, loosely rooted in the 
mud or firmly fixed in gravel. Leaves in a rosette; petiole (2.5 — )10 — 15 cm 
(depending on water depth), dark green, sheathed in the lower part for 
nearly one fifth of its length; leaf blade ovate, 4.5 — 8 cm long and 2 — 4 cm 
wide, base rounded, margin flat to slightly undulate, apex acute, upper 
surface dark green, sometimes with dark purple transverse markings, or 
submerged leaves brownish green; pale green or sometimes tinged reddish 
beneath. 

Peduncle 2 — 4 cm long. Spathe 10 — 15 cm long; limb of the spathe 
4 — 5 cm long and c. 1.5 cm wide, apex long caudate, base auriculate and 
margins reaching each other, upper surface pale yellow with distinct dark 
red irregular protuberances; collar distinct with a rim and forming a narrow 
opening to the tube; tube 3 — 6 cm long and c. 0.5 cm in diameter, outside 
and inside white: kettle 2 — 2.5 cm long, and c. 0.7 cm in diameter, outside 
white, inside white without purple markings. Spadix usually with 6 female 
flowers at the base, these pale green; stigmas white, more or less elliptic; 
olfactory bodies small, yellow; naked axis between male and female flowers 
c. 0.5 cm long, white; male part cylindrical, yellow, male flowers 50 - 60, 
each male flower usually consisting of two stamens; appendix cone shaped, 
c. 2 mm long, white: flap above the male flowers white. Fruit and seeds not 
known. 

Emersed cultivated plants much smaller; leaves c. 10 cm long; 
peduncle c. 1 cm long; spathe c. 6 cm long. 



12 



Gard. Bull. Singapore 53 1 2001) 




la. Dammed pools on the Jungle Fall Stream 
in the Bukit Timah Nature Reserve, 
Singapore. 



photo Bastmeijer 




lc. Cultivated plant of Cryptocoryne 
utimahensis. 

photo van Wijngaarden 



Figure 1. A new Cryptocoryne hybrid from 




lb. Limb of the spathe of Cryptocoryne 
xtimahensis. 

photo Bastmeijer 




Id. Opened kettle showing the female 
(bottom) and male flowers (top). 



photo van Wijngaarden 

Bukit Timah Nature Reserve, Singapore 



Cryptocoryne Xtimahensis from Singapore 



13 



Chromosome number: 2n = 54. Pollen fertility < 10%. Vouchers at 
Copenhagen (C). 

Other collections examined: All from Bukit Timah in Jungle Fall Valley - 
the type and J. Bastmeijer 807 (L), P. Blanc 91-1 (SING), K.S. Chua 333 
(SINIU D. Nicolson 1367 (US) and /. Sinclair 5200 (E). 

Discussion 

While plants identified as Cryptocoryne griffithii are known from several 
localities from Singapore, this new hybrid has only been collected from 
Bukit Timah. (There is just one collection. Ridley s.n. 1892 (SING), from 
Bukit Timah that may be C. griffithii). All collections of C. xtimahensis are 
from the Jungle Fall Valley, where the plant grows in two pools formed 
up- and down-stream by an artificial dam in the tiny stream (Fig. la). 
Constructed during the Japanese Occupation, it is currently silted up. The 
hybrid appears to be restricted to these pools. 

The first systematic analyses of pollen fertility and chromosome 
numbers in Cryptocoryne were made by Jacobsen (1977) and Arends et 
o/.(1982). With pollen fertility near zero for this plant, we are no doubt 
dealing with a hybrid. In addition, compared with C. griffithii in Singapore, 
for which half the herbarium specimens were collected in fruit, none of the 
herbarium specimens of this Bukit Timah population have fruits nor have 
fruits been observed in the field. 

The hybrid status is not. however, unique for Cryptocoryne. The 
popular aquarium species C. xwillisii Reitz from Sri Lanka forms a complex 
of hybrids in which C. parva De Wit is one of the parents, the other parent 
may be C. beckettii Trimen. C. walkeri Schott or C. undulata Wendt 
(Jacobsen 1981a.b). Another example is found in Tasek Bera. Peninsular 
Malaysia, where a very big population of C. purpurea grows (often 
erroneously ascribed to C. griffithii). which also has a pollen fertility of 
about zero and is today regarded as a hybrid between C. cordata and C 
griffithii (Jacobsen 1987). 

In classifying Cryptocoryne species, the main characters used are 
those of the limb of the spathe. C. xtimahensis has large, irregular, red 
protuberances on the limb of the spathe. the collar is very pronounced and 
the base of the limb is cordate (Fig. lb). These characters are distinct from 
those of C. nurii, where the limb of the spathe is rather elongated and has 
a yellow background, characters shared by C. cordata. 

In addition, the leaves of C. xtimahensis are narrowly ovate with a 
rounded to cordate base and an acute tip as are those of C. cordata. The 



14 



Card. Bull. Singapore 53 (2001) 



transverse markings on the upper leaf surface of most plants of C. 
xtimahensis (Fig. lc) are also a character of C. nurii. 

The combination of these characters makes C. griffithii less likely as 
one of the parents. The limb of C. griffithii is verrucose with small, regular 
bumps, although the collar is very pronounced too. However, the leaves of 
C. griffithii are broadly ovate. Both C. nurii and C. cordata grow in the 
southern part of Johore, Peninsular Malaysia but have never been recorded 
from Singapore. An artificial cross between these two species would be 
very interesting but would be very difficult to perform, because neither of 
the species is easy to cultivate. DNA analysis should be able to elucidate 
the exact parentage. 

Base chromosome numbers in Cryptocoryne are x = 10, 11, 14, 15, 17 
and 18. Polyploidy is rather common in Cryptocoryne and even rather long 
polyploid series exist (Arends et al., 1982). The chromosome number of 
this new hybrid, 2n = 54, is remarkable because it could indicate that this 
plant is of triploid origin with base number 18. This number is known only 
from three Sri Lankan species, the C. crispatula-group from mainland Asia, 
and the tidal species C. lingua Engl, from Sarawak. All known species 
from the Malay Peninsula have base number 17, except for the distinct C. 
longicauda Becc. ex Engl, (also known from Sumatra and Borneo), which 
has base number 15 and the tidal species C. ciliata, which has base number 
11. The number 2n=54 may be explained as originating from a triploid 
number with 2n = 51 after which a second generation of aneuploid 
segregation may have occurred (0rgaard et al. 1995). 

The Future of Cryptocoryne Habitats in Singapore 

The status of the Bukit Timah Nature Reserve provides a safe habitat for 
this plant. However, C. xtimahensis occupies a very restricted area c. 10 m 
long and 3 m wide. It is therefore extremely vulnerable to any hydrological 
changes to this one particular stream. C. griffithii seems to be rather common 
in the reservoirs (Chan, 2000), and provided there is no change in land use 
will not be threatened. 

C. ciliata, the other native Cryptocoryne of Singapore, is said to 
have been re-introduced with material originating from southern Johore. 

On the other hand, several species of Cryptocoryne have been 
introduced in the past, for example C. wendtii De Wit, C xwillisii Reitz 
from Sri Lanka, and C. lingua from Sarawak (Chan, 2000). They probably 
'escaped' by accident from the numerous nurseries, which produced - from 
the 1960's onwards - large quantities of these popular plants for aquascaping, 
mainly for export. It will be interesting to investigate the flora of Singapore 
in this respect. 



Cryptocoryne xlimahensis from Singapore 



15 



Acknowledgements 

The National Parks Board of Singapore is acknowledged for permission to 
collect in the Bukit Timah Nature Reserve and to J. Lai and S. Saifuddin 
for showing JDB the population. The aquatic plant firms Tropica 
(Denmark) and Oriental (Singapore) were of great help during JDB's stay 
in Singapore. Gitte Petersen (Copenhagen. Denmark) made the 
chromosome counts. Dr. H. Roessler (Munich. Germany) translated the 
diagnosis into Latin. Piet van Wijngaarden cultivated the plants in the 
Netherlands and was able to flower them after a couple of months. Niels 
Jacobsen (Agricultural University. Copenhagen. Denmark) and Josef 
Bogner (Botanical Garden. Munich. Germany) are acknowledged for their 
great support in discussing taxonomic problems in Cryptocoryne, and for 
comments on the manuscript. 

References 

Arends. J.C.. J.D. Bastmeijer and N. Jacobsen. 1982. Chromosome numbers 
and taxonomy in Cryptocoryne (Araceae).II. Nordic Journal of Botany. 
2: 453—463. 

Bastmeijer. J.D. 2000. The Crypts Pages, http://users.bart.nl/~cr> pis 
index.html 

Chan. S.Y. 2000. The Freshwater Macrophytes of Singapore, http:// 
molluscan.com/macrophytes 

Engler. A. 1920. Araceae - Aroideae Cryptocoryne. Das Pflanzenreich. 
IV.23.F: 232—249. 

Foulis. L. et al. (eds.) 1998. Botanica. 2 nd ed. Periplus Editions. Singapore. 

Hoedeman. J.J. 1948. Het geslacht Cryptocoryne Fischer, 1928. Het 
Aquarium 19(3): 61 — 64. 

Hooker. J.D. 1900. Cryptocoryne griffithii. Botanical Magazine 126, pi. 
7719. 

Jacobsen. N. 1977. Chromosome numbers and taxonomy in Cryptocoryne 
(Araceae). Botanisher Notiser. 130: 71-87. 

Jacobsen, N. 1981a. Cryptocoryne undulata Wendt und Bemerkungen zu 
anderen Arten. Aqua-Planta. 6(2): 31 — 38. 



16 



Card. Bull. Singapore 53 (2001) 



Jacobsen, N. 1981b. Cryptocoryne unditlata Wendt und Bemerkungen zu 
anderen Arten - Ein Nachtrag. Aqua-Planta. 6(4): 92 — 94. 

Jacobsen, N. 1982. Cryptocorynen. Kernen Verlag. Stuttgart, Germany. 

Jacobsen, N. 1987. Cryptocoryne purpurea Ridley. Aqua-Planta 12(2): 61 — 
62. 

Jacobsen, N. and J. Bogner. 1986. Die Cryptocorynen der Malaiischen 
Halbinsel (l.Teil). Aqua-Planta. 11(4): 135—139. 

Jacobsen, N. and J. Bogner. 1987. Die Cryptocorynen der Malaiischen 
Halbinsel (2.Teil). Aqua-Planta. 12(1): 13—20. 

Ong, Y.W. 1999. Private communication. Singapore. 

0rgaard, M., N. Jacobsen and J.S. Heslop-Harrison. 1995. Molecular 
cytogenetics in the genus Crocus L. In: P.E. Brandham and M.D. Benneth 
(eds.), Kew Chromosome Conference IV, pp. 291-299. Royal Botanic 
Gardens, Kew, U.K. 

Oskam, H.C. and J.D. van Ramshorst. 1954. Cryptocoryne griffithii Schott. 
Het Aquarium. 24(7): 146—149. 

Ridley, H.N. 1904. New Malayan Plants. Journal of the Straits Branch of 
the Royal Asiatic Society 41: 44 — 45. 

Ridley, H.N. 1907. Cryptocoryne. Materials for a Flora of the Malayan 
Peninsula. Pp. 3-6. 

Ridley, H.N. 1925. Cryptocoryne. The Flora of the Malay Peninsula. 5: 

86—88. 

Schott, H.W. 1856. Aroideae. Synopsis Aroidearum. 1: 1 — 3. 

Tan, H.T.W. 1995. A Guide to the Threatened Plants of Singapore. Singapore 
Science Centre, Singapore. 

Tan, H.T.W., I.M. Turner, Y.C. Wee and K.S. Chua. 1994. Plants in the 
Singapore Red List. Singapore Scientist. 72: 26 — 32. 

Wendt, A. 1952-1958. Die Gattung Cryptocoryne Fischer. Die 
Aquarienpflanzen in Wort und Bild. Stuttgart, Germany. 

Wit, H.C.D. de. 1951. Cryptocoryne cordata, Cryptocoryne griffithii. Het 
Aquarium. 22(6): 128—129. 



Cryptocoryne xiimahensis from Singapore 



17 



Wit. H.C.D. de. 1958a. Cryptocoryne griffithii. Fishkeeping. January: 138. 

Wit. H.C.D. de. 1958b. Cryptocoryne griffithii and its allies. Fishkeeping. 
February: 230. 

Wit. H.C.D. de. 1961. Het genus Cryptocoryne (17). Het Aquarium. 31(10): 
232—233. 

Wit, H.C.D. de. 1971. Aquarienpflanzen. Ulmer. Stuttgart. Germany. 
Wit. H.C.D. de. 1990. Aquarienpflanzen. 2 nd ed. Ulmer. Stuttgart. Germnay. 



Gardens' Bulletin Singapore 53 (2001) 19-74. 



The Genus Rhaphidophora Hassk. (Araceae- 
Monsteroideae-Monstereae) in Borneo 



P.C. Boyce 



Royal Botanic Gardens. Kew. Richmond. Surrey. TW9 3AE. U.K. 



Abstract 



An alpha-taxonomic account of Bornean Rhaphidophora is presented as a precursor to the 
forthcoming Flora Malesiana Araceae treatment. Thirteen species are recognized, none 
novel. Rhaphidophora elliptica Ridl. 1905 (non 1908 = R. elliptifolia Merr.) and R. 
megasperma Engi., treated respectively as synonyms of R. montana (Blume) Schott and R. 
puberula Engl, by Boyce (1999). are resurrected as Bornean endemics. New synonyms are: 
Scindapsus havilandii Ridl.. Rhaphidophora jacidiforrnis Alderw.. R. subfalcata M. Hotta 
into R. megasperma. and R. nigrescens Ridl. into R. sylvestris (Blume) Engl. A dichotomous 
key to species is provided. All species are illustrated. 



Rhaphidophora Hassk. (including Afrorhaphidophora Engl.; c. 3 species in 
tropical Africa) comprises c. 100 species of small to large, occasionally 
enormous, root-climbing lianes (sensu Schimper, 1903). rarely rheophytes, 
distributed from tropical West Africa eastwards to the western Pacific, 
north to southern Japan (Ryukyu Islands) and south to Northern Australia. 
Rhaphidophora is one of the largest aroid genera represented in tropical 
Asia and has several nodes of diversity: the Himalaya (SE Nepal to NE 
Vietnam, roughly 17° — 23° N), West Malesia (including southernmost 
peninsular Thailand), the Philippines, and East Malesia. 

This is the fifth in a series of papers intended to present a complete 



Contents 



Introduction 

Geography and endemism 

Rhaphidophora 

Key to adult flowering Rhaphidophora in Borneo 

The species 

Acknowledgements 

References 

Index exsiccatorum 



19 
20 
21 
24 
26 
72 
72 
73 



Introduction 



20 



Gard. Bull Singapore 53 12001) 



alpha-taxonomy of the genus Rhaphidophora in Asia. Accounts for 
Peninsular Malaysia and Singapore, the southern and western Indonesian 
archipelago and the Philippines have been published recently (Boyce, 1999, 
2000a, 2000b) together with a partial account for New Guinea (Boyce & 
Bogner, 2000) and an account for Papuasia and the tropical Western Pacific 
appears elsewhere in this journal (Boyce, 2001). Accounts for the Himalaya, 
and Thailand and Indochina are being prepared and will be published 
separately. All morphological terms employed follow Stearn (1992). 

The last complete revision of Rhaphidophora was that of Engler & 
Krause (1908). A summary of the taxonomic and nomenclatural history of 
Rhaphidophora was presented in Boyce (1999) and a partial infrageneric 
classification with an overview of informal morpho-taxonomic groups was 
proposed by Boyce (2000a). Boyce (1999) provides a detailed discussion of 
structure, generic limits, together with keys to the genera of monsteroid 
and anadendroid aroids in Malesia. 

Synonymy cited is for the species, not for the review area. The reason 
for this is that synonymic names based on types from outside the review 
area are frequently applied to specimens in local herbaria. 

Geography and Endemism 

Borneo has a relatively poor Rhaphidophora flora, with comparatively few 
species recorded, although how much this is a reflection of inadequate 
collecting is not clear. However, of the 13 species recognized for the island 
five are endemic (compared with 2/15 in Peninsular Malaysia; 4/15 in 
Sumatera) and thus Borneo has, with the exception of the Philippine islands 
(7/11), the highest level of endemism in Sunda. 

Two of the five Bornean endemics, R. elliptica Ridl. and R. ellipti folia, 
belong to the Indo-Malesian Hongkongensis Group (see Boyce, 2000a). 
Both Bornean endemics are morphologically close to west Malesian species 
(R. elliptica to R. montana (Blume) Schott; R. ellipti folia to R. maingayi 
Hook./.) and are presumably endemic segregates. Another species from 
this group, the widespread R. sylvestris (Blume) Engl. (Sumatera, Peninsular 
Malaysia, Thailand, east to Borneo) is widespread and common in Borneo. 
The other Bornean endemics are R. latevaginata M. Hotta, R. megasperma 
Engl, and R. cylindrosperma Engl. & K. Krause. 

Rhaphidophora latevaginata M. Hotta is a neotenic shingling species 
that might conceivably have arisen from a species with a pronounced 
heteroblastic development, e.g., R. korthalsii Schott (see Boyce, 2000a). 

Rhaphidophora megasperma is of uncertain affinity in Sundaic Asia 
but has morphological similarities, e.g., clustered inflorescences subtended 



Rhaphidophora in Borneo 



21 



by chartaceous cataphylls. to certain New Guinea species (e.g., R. versteegii 
Engl.) and is tentatively included in the Hollrungii Group (note that this 
group had its name changed in Boyce. 2001). 

Rhaphidophora cylindrosperma is another morphologically isolated 
species in Sunda but with similar species (e.g.. R. schlechteri K. Krause) 
east of Wallace s Line and belongs to the Neoguineensis Group. It is 
interesting that two of the Borneo endemic Rhaphidophora appear to be 
morphologically closer to species in Wallacean New Guinea than to any 
Sundaic species. 

RHAPHIDOPHORA 

Rhaphidophora Hassk.. Flora 25 (2) Beibl. 1 (1842) 11; Schott, Gen. Aroid. 
(1858) 77 & Prodr. Syst. Aroid. (1860) 377—388; Miquel, Ann. Mus. Bot. 
Lugd.-Bat. 3 (1867) 81—82; Engl, in A. & C. DC., Monogr. Phan. 2 (1879) 
238—248; Engl, in Beccari. Malesia 1 (1882) 266—272, Tab. xix 6—9. xx 
1—5; Benth. & Hook, f.. Gen. PI. 3(2) (1883) 993 - 993; Engl. & Prantl, 
Nat. Pflanzenfam. T. 2. Ab. 3 (1889) 119—120; Engl. & Prantl. Nat. 
Pflanzenfam. Nachtr. 1 (1897) 58: Ridl.. J. Straits Branch Roy. Asiat. Soc. 
44: 185—187 (1905); Engl. & K. Krause in Engl., Pflanzenr. 37 (IV.23B) 
(1908) 17—53: Engl. & Prantl, Nat. Pflanzenfam. Nachtr. 3 (1908) 29; 
Merrill. J. Straits Branch Roy. Asiat. Soc. special number (1921) 88 — 90; 
Schott, Icones aroideae et reliquiae (IDC Microfilm) (1983) fiche nos. 
28—31. 121: Mayo et al, Genera Araceae (1997) 118—121. PI. 14, 109 D. 

— Scindapsus Schott subgen. Rhaphidophora (Hassk.) Miq.. Flora Ned. 
Indie 3 (1856) 185. — Type: Rhaphidophora lacera Hasskarl, nom. Meg. 
pro. Pothos pertusus Roxb. [= Rhaphidophora pertusa (Roxb.) Schott] 

Scindapsus Schott subgen. Pothopsis Miq.. Flora Ned. Indie 3 (1856) 187. 

— Type: Scindapsus sylvestris (Blume) Kunth [= Rhaphidophora sylvestris 
(Blume) Engl.] 

[Raphidophora Hassk.. Cat. Hort. Bogor. (1844) 58, orth. var.] 

Medium-sized to very large, occasionally enormous, slender to robust, 
leptocaul or pachycaul, homeophyllous or heterophyllous, rarely neotenic 
[e.g.. some populations of R. beccarii (Engl.) Engl.], root-climbing lianes, 
very seldom clustering and rheophytic (e.g.. R. beccarii) and then always 
with a creeping juvenile stage; cut surfaces producing clear, odourless sticky 
juice either drying ± invisible or coagulating into yellowish, translucent 
jelly and eventually hardening to a brittle amber-like mass; seedling stage 



22 



Card. Bull. Singapore 53 (2001) 



mostly not observed but where known either leafy at germination and 
skototropic (see Strong & Ray, 1975) by an alternating series of congested 
leafy and elongated leafless shoots (e.g., R. angustata Schott) or germinating 
to give rise to a non-skototropic shingling juvenile shoot (e.g., R. korthalsii); 
pre-adult plants often forming modest to extensive terrestrial colonies of 
varying morphological and physiological form (descriptive generalisations 
are nearly impossible), largest terrestrial colonies generally occurring in 
places of less than optimum adult growth potential (e.g., depauperate tree 
canopy, dry, exposed sites); adult shoot architecture broadly divisible into 
three types: i. physiognomically unbranched clinging non-flowering stems 
rooting along their entire length giving rise to variously elaborated free 
lateral flowering stems (e.g., R. lobbii Schott, R. puberula Engl.), or ii. all 
stems physiognomically unbranched clinging and flowering (e.g., R. 
korthalsii), or hi. physiognomically unbranched leader and lateral stems 
clinging but only lateral stems flowering (e.g., R. foraminifera): stems with 
internodes of various lengths separated by variously prominent leaf scars, 
smooth or asperous or densely pubescent to ramentose (the last not in the 
review area), older stems subwoody or somewhat corky or with distinctive 
matt to sublustrous pale brown papery epidermis, with or without variously 
textured prophyll, cataphyll and petiolar sheath fibre either at the tips or 
along the newer sections, rarely with cataphylls and prophylls deliquescing 
to black mucilage later drying to leave fragmentary parchment-like remains 
on petioles, developing laminas and inflorescences; flagellate foraging stems 
occurring in some species, often exceedingly long, reaching the ground 
then rooting, variously foraging and climbing again; clasping roots sparsely 
to densely arising from the nodes and internodes, clinging to substrate; 
feeding roots rare to abundant, smooth, pubescent or prominently scaly, 
later often becoming woody, clinging to substrate or free; leaves distichous 
or weakly spiralled, evenly distributed or scattered or clustered distally; 
cataphylls and prophylls subcoriaceous to membranous, either soon drying 
and falling or degrading or deliquescing to variously textured sheaths and 
fibres, these where present variously clothing upper stem before eventually 
decaying and falling; petiole canaliculate to weakly carinate, smooth or 
pubescent, with variously prominent apical and basal genicula; petiolar 
sheath prominent, nearly reaching to overtop the geniculum, occasionally 
one side greatly expanded and auriculate. especially in juvenile plants, at 
first membranous to coriaceous, soon completely or along the margins 
drying chartaceous, sometimes degrading to untidy variously netted or 
simple fibres and later variously falling to leave a scar or disintegrating 
marginally or completely; lamina submembranous to stiffly chartaceous or 
coriaceous, lanceolate or oblong, ± oblique, base decurrent to unequal or 
cordate, apex acute to acuminate, entire to regularly pinnatifid or 



Rhaphidophora in Borneo 



23 



perforated, if pinnate then divisions pinnatifid to pinnatisect (Stearn. 1992: 
324), midrib often ± naked between segments, lamina occasionally with 
small to well developed perforations adjacent to the midrib and primary 
veins, these sometimes extending to lamina margin (fenestrations then 
occasionally additional to fully developed pinnae), rarely abaxially 
pubescent when expanding, rarely strongly concolorous at maturity: midrib 
usually prominent raised abaxially and prominently sunken, sometime flush, 
rarely slightly raised adaxially: primary venation ± pinnate: interprimaries 
mostly present, subparallel to primaries and sometimes indistinguishable 
from them (e.g.. R. monticola - Philippines) but usually less prominent and 
often drying paler, usually glabrous, occasionally pubescent with domatia 
in the axils of the primary and secondary veins: secondary venation striate 
(e.g.. R. monticola — Philippines) to reticulate (e.g.. R. korthalsii). variously 
prominent, often very difficult to distinguish from primary venation (e.g.. 
R. angustata); tertiary venation where visible reticulate to tessellate: 
inflorescences solitary to several together, first inflorescence subtended by 
a (usually fully developed) foliage leaf and/or a very soon disintegrating 
cataphyll. subsequent inflorescences usually each subtended by a prophyll 
and cataphyll. more rarely by a prophyll and partially to almost fully formed 
foliage leaf, inflorescences at male anthesis naked by disintegration of 
subtending cataphyll or partially to almost completely obscured by netted 
and sheet-like fibres; peduncle terete to laterally compressed: spathe ovate 
to narrowly or broadly canoe-shaped, stoutly to rather weakly beaked, 
barely gaping to opening almost flat at male anthesis and then usually 
deciduous before male anthesis is complete, occasionally persisting into 
the early stages of infructescence development (e.g.. R. angustata). rarely 
drying and persistent (e.g., R. megasperma). stiff to rather softly or stoutly 
coriaceous, dirty-white, greenish, cream or yellow; spadix subglobose to 
clavate-cylindrical. cylindrical or fusiform, sessile or stipitate. often obliquely 
inserted on peduncle, tapering towards the apex; flowers bisexual, naked: 
ovary 1- to partially 2-locular. lower part ± bilaterally compressed, upper 
part ± cylindrical and variously angled, most often rhombohexagonal. those 
upper- and lower-most on the spadix often sterile and bereft of stigma, 
those uppermost frequently either scattered or partially fused to each other 
and forming a rudimentary appendix: ovules few to many, anatropous. 
funicle long, placentae parietal to basal, sometimes ± subaxile. partial septa 
variably intrusive: stylar region well developed, usually broader than ovary, 
usually truncate apically. rarely elongate-conic: stigma sticky at female 
male anthesis. punctiform. broadly elliptic or oblong, orientation 
circumferential or longitudinal: stamens A — 6; filaments strap-shaped: anthers 
usually prominently exserted from between ovaries at male anthesis. rarely 
not exserted and pollen extruded from between ovaries, thecae dehiscing 



24 



Card. Bull. Singapore 53 (2001) 



by a longitudinal slit; infructescence with stylar regions greatly enlarged, 
transversely dehiscent, the abscission developing at the base of the enlarged 
to massive stylar region and this falling to expose the ovary cavity with the 
many seeds embedded in variously coloured sticky pulp; seeds oblong, 
testa thin, smooth, embryo axile, straight, endosperm copious; pollen 
dicolpate, extended monosulcate to fully zonate, ellipsoid or hamburger- 
shaped, medium-sized (mean 33 urn, range 24 — 55 urn) (Mayo et al, 1997), 
exine foveolate, subreticulate, rugulate, fossulate, scabrate, retiscabrate, 
verrucate, or psilate; chromosomes 2n = 60, 120 (42, 54, 56) (Mayo et al, 
1997). 

Distribution: About 100 species from tropical Africa, South and South 
East Asia, Australia and the Pacific with extensions into the subtropical 
Himalaya, southern China and the southernmost islands of Japan. 

Habitat: Lianescent bole-climbers, lithophytes, rarely rheophytes, usually 
in well drained subtropical and tropical wet, humid, or seasonally 
moderately dry primary and established secondary evergreen forest at low 
to mid-montane elevations. 

Etymology: Greek rhaphis, rhaphidos (needle) and phero (I bear); refers 
to the macroscopic (to 1 cm long), needle-like unicellular trichosclereids 
present in tissues. 



Key to Adult Flowering Rhaphidophora in Borneo 

la. Mature leaf lamina pinnately divided 2 

lb. Mature leaf lamina entire, without or without perforations, but never 
pinnately divided 3 



2a. Flowering plants usually rheophytic. rarely low-climbing on trees 
beside torrential streams. Plants always associated with sandy or 
rocky forest streams 1. R. beccarii 

2b. Flowering plants high-climbing on forest trees. Plants not specifically 
associated with watercourses 7. R. korthalsii 



3a. Geniculum and abaxial surface of lamina pubescent 4 

3b. Geniculum and abaxial surface of lamina glabrous 5 

4a. Flowering on clinging stems. Leaves of mature plants extensively 
perforated. Active shoot tips with black mucilage 



Rhaphidophora in Borneo 



25 



6. R. foraminifera 

4b. Flowering on free lateral stems. Leaves of mature plants lacking or 
with only with scattered, perforations. Active shoot tips lacking black 
mucilage 12. R. puberula 

5a. Leaves always shingling, even in flowering individuals; leaf laminas 
stiffly coriaceous, broadly oblong-ovate-elliptic, 8 — 48 x 6.5 — 20.5 
cm, bright green, slightly to markedly glaucous, base truncate-cordate 

to broadly cuneate. Flowering on clinging shoots 

8. R. latevaginata 

5b. Leaves spreading in adult and flowering individuals; leaf laminas 



variously coloured but never glaucous. Flowering on free or clinging 
shoots 6 

6a. Stems scabrid to asperous. Spathe exterior minutely puberulent .... 

9. R. lobbii 

6b. Stems smooth. Spathe exterior glabrous 7 



7a. Clinging stems square or rectangular in cross-section; tips of active 

stems with netted prophyll, cataphyll and petiolar sheath fibre 

4. R. elliptica 

7b. Clinging stems variously shaped in cross-section, including square 



or rectangular; tips of active stems without fibres 8 

8a. Spadix stipitate; leaf lamina thickly coriaceous.... 2. R. conocephala 
8b. Spadix sessile; leaf lamina variously textured but never thickly 
coriaceous 9 



9a. Inflorescences two, three or more together, each subtended by a 
prominent chartaceous prophyll and one or more chartaceous 
cataphyll; leaf lamina oblong-lanceolate or oblong-elliptic, entire to 
slightly perforated, glabrous 10. R. megasperma 

9b. Inflorescences almost always solitary, if two together then 
inflorescences not subtended by prominent chartaceous prophylls 
and cataphylls; leaf lamina variously shaped and almost never 
perforated, if perforated then apical geniculum and abaxial surface 
of lamina pubescent 10 



10a. 
10b. 



Clinging stems rectangular in cross-section 

Clinging stems more-or-less terete in cross-section 



11 
12 



26 



Card. Bull. Singapore 53 (2001) 



11a. Leaf lamina lanceolate-elliptic to falcate-lanceolate, 4.5 — 32 x 1.75 — 
8.5 cm. Spadix cylindric to weakly clavate-cylindrical, 3 — 6 cm long 
13. R. sylvestris 

lib. Leaf lamina narrowly elliptic to narrowly elliptic-oblong, 20 — 29 x 
4.5 — 7 cm. Spadix cylindrical, c. 5 x 1 cm 5. R. elliptifolia 

12a. Leaf lamina narrowly falcate-elliptic to falcate-lanceolate or falcate- 
oblanceolate, 2.5 — 16 x 1.2 — 3 cm, drying uniformly pale straw- 
coloured. Spadix slender cylindrical, 2.5 — 7 cm long ... 11. R. minor 

12b. Leaf lamina subfalcate-lanceolate or oblong-elliptic, oblique, 10 — 
19.5 x 2.5 — 6.5 cm, drying very dark brown. Spadix cylindrical-obtuse, 
3 — 10 x 1 cm (fruiting specimen only) 3. R. cylindrosperma 

The Species 
1. Rhaphidophora beccarii (Engl.) Engl. 

Rhaphidophora beccarii (Engl.) Engl, in Bot. Jahrb. Syst. 1 (1881) 181 & in 
Beccari, Malesia 1 (1882) 270, Tab. xix 6—9; Engl. & K. Krause in Engl., 
Pflanzenr. 37 (IV.23B) (1908) 46; Alderw., Bull. Jard. Bot. Buitenzorg III, 
1 (1920) 382; Merrill, J. Straits Branch Roy. Asiat. Soc, special number 
(1921) 88; Alderw., Bull. Jard. Bot. Buitenzorg III, 4 (1922) 341. — 
Epipremnum beccarii Engl.. Bull. Soc. Tosc. Ortic. 4: 268 (1879). — Type: 
Malaysia, Sarawak, Kuching, Oct. 1865, Beccari PB 833 (cited as '832' by 
Engler. 1879) (FI, holo). 

Rhaphidophora borneensis Engl, Araceae exsicc. et illustr. n. 195 & in Bot. 
Jahrb. Syst. 7, Beibl. 15 (1886) 1. — Type: Indonesia, Kalimantan, Mindai 
to Pramassan, 19 June 1882, H. Grabowski s.n. (Bf, BM). 

Rhaphidophora fluminea Ridl., J. Straits Branch Roy. Asiat. Soc. 44 (1905) 
186; Engl. & K. Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 37; Merrill, 
J. Straits Branch Roy. Asiat. Soc, special number (1921) 89. — Type: 
Malaysia, Sabah, Bongaya, Dec. 1897, Ridley s.n. (SING, holo). 

Figure 1 

Small to medium-sized, heterophyllous, sometimes homeophyllous, creeping 
rheophyte, very rarely short liane, to 75 cm; seedling stage a non-skototropic 
shingling shoot, soon becoming spreading-leafy; pre-adidt plant initially 
with + appressed shingle-leaves, later with leaves erect or spreading and at 
this stage plants resembling adults in all but overall size and leaf division; 



Rhaphidophora in Borneo 



27 




Figure L Rhaphidophora beccarii (Engl.) Engl. 

A. habit x '/.,: B. leaf lamina x 7 4 ; C. leaf lamina x l / 4 ; D. venation detail x 4: E. inflorescence, 
spathe fallen x L; F. spadix detail, post-female anthesis. pre-male anthesis x 8. A. E & F from 
Stone 9637: B & D from Ridley s.n.: C from Kiew 1982. 



28 



Card. Bull. Singapore 53 (2001) 



adult shoots clinging and flowering but main axis comprised of longer 
modules than axillary axes; stems smooth, mid- to dark green, with very 
sparse petiolar sheath fibre, internodes 1 — 7 x c. 1 cm, separated by variously 
prominent slightly oblique leaf scars; flagellate foraging stems absent; 
clasping roots very densely arising from the nodes and internodes of clinging 
stems, pubescent; feeding roots clinging, densely scaly; leaves distichous, 
appressed and shingling to erect or spreading and scattered on pre-adult 
shoots, tending to become distally clustered on adult shoots; cataphylls and 
prophylls membranous, soon drying and falling; petiole narrowly canaliculate 
to carinate, 8 — 31 x 0.3 — 1.5 cm, smooth, apical and basal genicula 
prominent; petiolar sheath prominent, extending to the apical geniculum, 
variably persistent and mostly degrading into semi-persistent weak fibres; 
lamina entire in seedling and pre-adult individuals, entire, pinnatipartite 
or pinnatisect in adult plants, narrowly lanceolate to oblong-elliptic, slightly 
oblique, 21 — 51 x 2 — 23 cm, subcoriaceous to slightly fleshy, base decurrent, 
apex acuminate with a moderately prominent apicule; midrib prominently 
raised abaxially, sunken adaxially; primary venation pinnate, raised abaxially, 
slightly impressed adaxially; interprimaries subparallel to primaries, slightly 
raised abaxially, ± flush adaxially, often forming a weak reticulum; secondary 
venation prominently reticulate, slightly raised: tertiary venation a network 
of broadly spaced tessellate veins arising at c. 90° from the midrib and 
crossing the primaries and interprimaries; inflorescences one to three 
together, subtended by a prominent cataphyll degrading to fibres before 
male anthesis; peduncle terete, 8 — 12 x 0.2 — 0.4 cm; spathe narrowly canoe- 
shaped, stoutly beaked, 6.5 — 7 x 1 — 1.5 cm, stiffly fleshy, greenish to dull 
white, soon falling at male anthesis; spadix cylindrical, sessile, inserted 
perpendicular to peduncle, 4.5 — 7 x 0.6 — 1 cm, dull whitish; stylar region 
rather well developed, mostly rhombohexagonal, c. 1 — 1.5 x 1 — 1.5 mm, 
truncate; stigma elliptic, longitudinally orientated, occasionally almost 
circular, c. 0.5 x 0.3 mm, often very prominent especially in dried material; 
anthers exserted at male anthesis; infructescence 7—9 x 1 — 2 cm, mid- 
green when ripe. 

Distribution: Southern Thailand, Peninsular Malaysia, Sumatera (including 
the Lingga Archipelago) and Borneo (Sarawak, Brunei Darussalam, Sabah, 
West and East Kalimantan). 

Habitat: Along rocky or sandy stream banks, often on rocks in stream or at 
waterfalls in primary to slightly disturbed or old secondary lowland forest, 
peatswamp forest, occasionally persisting briefly in logged over areas, 
occurring on a variety of substrates. Sea level to 300 m altitude. 



Rhaphidophora in Borneo 



29 



Notes: 1. One of possibly two obligate rheophytic species (the other is 
Rhaphidophora araea P.C. Boyce - Sumatera). R. beccarii is immediately 
recognizable by its adult growth form, creeping along watercourses or 
attached on rocks in the water, and by the usually pinnately divided leaf 
laminas. Occasionally plants are seen climbing up tree trunks on the bank 
of torrential streams. 

2. Although the pinnately divided leaf is typical of adult plants, entire 
leaved stenophyllous to lanceolate-leaved flowering plants are not rare. 
Such plants have been referred to as R. fluminea and occur either as pure 
stands or as mixed populations with the pinnate-leaved form. They are 
treated here as a neotenic manifestation of R. beccarii. 

3. Rhaphidophora borneensis. based on a Grabowski specimen from 
Kalimantan, is a broad-leaved form of typical (i.e.. divided-leaf) R. beccarii. 

Other Bornean specimens seen: SARAWAK. 1 Div.: Setapok F.R.. 6 miles south of Kuching. 
Bogner 1507 (K. M); Sg. Entabai. Bogner 1562 (M, US); Matang Family Park. Sg. Cina, 
Boyce 719 (K): Lundu. Brooke 8409 (L):13th Mile. Matang. Brooke 9459 (L): Lundu. G. 
Gadin. Clemens & Clemens 21934 (K): Matang F.R.. 10 miles west of Kuching. Nicolson 
1282 (US); Setapok F.R.. 6 miles south of Kuching, Nicolson 1343 (L, US); Matang. Ridley 
s.n. (BM): Padawan. G. Merubong. Ulu Sg. Sluba. Yii S51363 (K. L. SAR. US); 2" d Div.: 
Simanggang. Brooke 10729 (BM. L); 3' d Div.: Kapit District. S. Bena area. Burn 13001. 
12938 (E): 4' h Div.: Lambir N.P., Mile 18. Chai S39439 (K. KEP. L. SAR. US): 5' h Div.: 8 - 9 
miles from Limbang. Sg. Bakol. Bogner 1504 (M. US): 7 Div.: En route (survey highway) 
from Sg. Mah to Sg. Shinonok. Ulu Sg. Minah. Bintulu District. Hirano & Hotta 14082 
(KYO): Sg. Bejangung. a branch of the Sg. Anap. Bintulu District. Hirano & Hotta 1174 
(KYO): not located: Sg. Entabai. Bogner 1346 (K. M. US); Sg. Engkramut. Bogner 1393 
(M. US). BRUNEI DARUSSALAM. Belait: Labi, Kg. Teraja. path along Sg. Teraja, 
Boyce 245 (BRUN. K. L): Temburong: Sg. Temburong at Kuala Belalong. Boyce 396 
(BRUN. K. L): Sg. Temburong near Kuala Belalong. Jacobs s.n. (BRUN. K. L); Sg. 
Temburong. just downstream from Kuala Belalong. Wong 242 (BRUN. K. L). SABAH. 
Kudat: Ranau. Kg. Puas area. Amin et al. SAN 94681 (K. KEP. L. SAN. SAR. SING): 
Pantai Barat: Keningau. Ulu Sg. Pingas Pingas. Jimpin SAN 122024 (K, KEP, SAN): 
Keningau. Shang Lian logging area. Lanas. Krispinus SAN 118444 (K. SAN): Keningau. 
Camp C area, fiulan. Lantoh SAN 102053 (K. KEP. L. SAN): Keningau, Ulu Sg. Matud. 
Tangki SAN 119596 (K. L, SAN. SAR); Sandakan: Lamag. Ulu Sg. Lokan, Aban & Petrus 
SAN 90675 (K. SAN): Sg. Lokan. Amin et al. SAN 97483 (K. L. SAN. SAR): VJH 45 A 
Lungmanis, Dewol et al. SAN 118075 (K. KEP. L. SAN): Beluran, Sg. Baba Gibot SAN 
90040 (K, L. SAN): Labuk and Sugut. western side of Bt. Doji and pass way from Telupid 
to Ulu Karamuak. Kokawa & Hotta 471 (KYO. L); Telupid. Labuk and Sugut. along Sg. 
Meliau. foot of G. Tawai. Kokawa & Hotta 110 (KYO. L): Mile 45 Labuk road. Meijer 
SAN 44014 (K. L. SAN): KALIMANTAN. West Kalimantan: Selimban Kapuas. Main 
(sub. Polak) 2092 (L): Bt. Ubili. Nieuwenhuis 97 (BO): East Kalimantan: W. Koetai, No. 29 
near L. Liah Leng. Endert 3013 (BO, K, L); W. Koetai, No. 5 near Lahoem. Endert 1860 
(BO. K. L): Kelassar (Kelasen?). Hallier 1555 (BO): Pulau Nibung, W. Koetai, Sg. Loewai. 
near Padang Loewai. Posthitmus 2173 (BO): Samarinda. mouth of Batang Mahakam. Sg. 



30 



Card. Bull. Singapore 53 (2001 ) 



Mukun. near Sanga Sanga. Meijer 1136 (BO. L): No further data. Hallier 999 (BO): Batu 
Lesoena, Nieuwenhuis 219 (BO). 

2. Rhaphidophora conocephala Alderw. 

Rhaphidophora conocephala Alderw., Bull. Jard. Bot. Buitenzorg III, 1 
(1920) 384. — Type: Indonesia. Sumatera, North Sumatera, Sibolangit, 10 
May 1917, Lorzing 5137 (BO, holo; K, L, iso). 

Figure 2 

Large, moderately robust, semi-pachycaul homeophyllous liane to 15 m: 
seedling stage a non-skototropic shingling shoot; pre-adult plants forming 
small terrestrial colonies of appressed shingling shoots; adult shoot 
architecture comprised of elongated, clinging, physiognomically unbranched, 
leafy, non-flowering stems and moderately elaborated, free, moderately 
leafy, flowering stems; stems smooth, climbing stems rectangular in cross- 
section, free stems more or less terete to very weakly four-angled in cross- 
section, larger shoot systems pendent under their own weight, without 
prophyll, cataphyll and petiolar sheath fibre, internodes to 7 x 1.5 cm on 
clinging shoots, shorter and less stout on free shoots, separated by prominent 
oblique leaf scars, older stems woody; flagellate foraging stems absent; 
clasping roots densely arising from the nodes and internodes of clinging 
stems, densely pubescent; feeding roots rare, clinging, pubescent; leaves 
weakly spiralled on clinging shoots, slightly scattered-distichous on flowering 
shoots; cataphylls and prophylls membranous, very soon drying and falling: 
petiole shallowly canaliculate adaxially. 4 — 7 x 0.1 — 0.2 cm, smooth, with a 
slight apical and rather prominent basal geniculum; petiolar sheath 
prominent, extending to and encircling the apical geniculum, very soon 
drying and falling to leave a thin continuous scar from the petiole base, 
around the top of the apical geniculum and back to the base, occasionally 
newest leaves with parchment-like sheath remain briefly adherent: lamina 
entire, falcate-lanceolate to falcate-oblong or falcate-oblanceolate, 10 — 
29.5 x 1.5 — 7 cm, coriaceous, upper surfaces slightly glossy, lower surfaces 
less so, base minutely cordate to subovate to acute or briefly decurrent, 
apex subacute with a prominent apiculate apicule, margins slightly revolute 
in dried material: midrib raised abaxially, very slightly sunken adaxially; 
primary venation pinnate, raised abaxially and adaxially; interprimaries 
parallel to primaries, slightly raised abaxially and adaxially: secondary and 
tertiary venation slightly raised in dried specimens; inflorescence solitary, 
subtended either by a fully developed foliage leaf or by one or more 
subfoliar (i.e., developed petiole but atrophied lamina) cataphyll; peduncle 
slightly compressed-cylindric, 3 — 6 x 0.3 — 0.5 cm: spathe cigar-shaped. 



Rhaphidophora in Borneo 



31 




Figure 2. Rhaphidophora conocephala Alderw. 

A. flowering shoot x V : ; B. inflorescence, spathe fallen x 1V 2 ; C. spadix detail, female anthesis 
x 10; D. gynoecium, three quarter view x 6; E. spadix detail, post-male anthesis x 10; F. portion 
of pre-adult sterile shoot x '/,: G. leaf lamina x 1 : : H. venation detail x 3. A - E, G. & H from 
Nur SFN 7369; F from Lorzing 11750. 



32 



Gard. Bull. Singapore 53 (2001) 



stoutly short-beaked, 7 — 9.5 x 2 — 3.5 cm, thickly fleshy, exterior light yellow, 
interior darker, soon (?) falling at female anthesis; spadix cylindrical to 
slightly clavate, very shortly stipitate, light yellow, 4 — 5.3 x 1.2 — 1.5 cm; 
stipe c. 2 mm long; stylar region well developed, mostly rounded to 
rhombohexagonal, 1.2 — 1.3 x c. 1.2 mm, conical; stigma conspicuously raised- 
punctiform, c. 0.2 mm diam.; anthers slightly exserted at male anthesis; 
infructescence stoutly cylindrical, 6.5 — 7.5 x 1.8 — 2 cm. 

Distribution: Sumatera, Kalimantan (East) and into the Philippines 
(Palawan). 

Habitat: Damp primary and old secondary forest. 450 — 1000 m altitude. 

Note: Very close to and possibly indistinguishable from Rhaphidophora 
sylvestris but consistent in the conical style topped with a prominent, raised 
button-like stigma and the notably more coriaceous leaves. 

Other Bornean specimens seen: KALIMANTAN. East Kalimantan: Batu Penalong, 
Mahakam, Wiriadinata 850 (BO, L). 

3. Rhaphidophora cylindrosperma Engl. & K. Krause 

Rhaphidophora cylindrosperma Engl. & K. Krause, Pflanzenr. 37 (IV.23B) 
(1908) 28; Merrill, J. Straits Branch Roy. Asiat. Soc, special number (1921) 
88. — Type: Indonesia, northern West Kalimantan, between the greater 
and lesser arms of the S. Sambar, Hallier 1164 (BO, holo). 

Figure 3 

Moderate leptocaul homeophyllous liane to unknown ultimate height; 
seedling stage and pre-adult plants not observed; adult shoot architecture 
not fully observed, flowering stems, moderately branched and leafy, 
apparently pendent; stems smooth, terete, with cataphylls and prophylls 
soon falling, internodes 2 — 3.5 x 0.8 — 1.2 cm, separated by prominent 
straight to slightly oblique leaf scars; flagellate foraging stem not observed; 
roots not observed; leaves spiro-distichous; cataphylls and prophylls 
membranous, soon falling; petiole canaliculate, 5 — 14 x 0.2 — 0.4 cm, basal 
geniculum very large and prominent, apical geniculum less so; petiolar 
sheath narrow and not prominent, extending to apical geniculum, + short- 
persistent; lamina entire, subfalcate-lanceolate or oblong-elliptic, oblique, 
10 — 19.5 x 2.5 — 6.5 cm, membranous, base acute, apex acuminate; midrib 
prominently raised abaxially, slightly impressed adaxially; primary venation 
pinnate, prominently raised abaxially, impressed adaxially; interprimaries 



Rhaphidophora in Borneo 



33 




Figure 3. Rhaphidophora cvlindrosperma Engl. & K. Krause 

A. flowering shoot x '/,; B. flowering shoot x V : : C. leaf lamina x 'U: D. venation detail x 2: E. 
inflorescence, spathe fallen x 1;F. spadix detail, post-male anthesis x 6. A. C - D from Church 
et al 1573: B. E - F from Sidiyasa PBU 650. 



34 



Card. Bull. Singapore 53 (2001) 



subparallel to primaries, slightly less prominent; secondary venation 
reticulate, slightly raised abaxially; inflorescence solitary, subtended by a ± 
fully developed foliage leaf and one or more cataphylls; peduncle terete, 

2 — 4 x 0.3 — 0.4 cm; spathe slender, 9 — 11 x 0.5 cm, apically tapering and 
ultimately stout beaked; spadix cylindrical-obtuse, sessile, inserted ± on 
peduncle, 3 — 10 x 1 cm (fruiting specimen only); stylar region moderately 
developed, slightly rhombohexagonal, c. 1.8 mm diam., ± truncate; stigma 
punctiform, raised, c. 0.3 mm diam.; anthers not observed; infructescence 

3— 10 x 1—1.5 cm. 

Distribution: Endemic to Borneo. Sarawak (I s Division), Brunei Darussalam 
(Belait), Kalimantan (West, Central and East). 

Habitat: Lowland primary to lightly disturbed forest. 25 — 200 m altitude. 

Notes: I. Superficially similar to Rhaphidophora puberula but differing in 
leaves completely glabrous and never perforated, by the smaller, more 
slender spadix and in flowering on long, free shoots. 

2. Among the Sundaic Asian species, it is not immediately apparent to 
what R. cylindrosperma is related. The superficial similarity to species in 
the Hookeri Group (e.g., R. puberula and R. foraminifera) is not borne out 
by closer examination. The growth form with inflorescences often carried 
at the tips of short shoots arising from much longer pendent stems is 
reminiscent of R. lobbii, but R. cylindrosperma differs in more coriaceous 
leaves, different leaf venation and in having a longer, externally glabrous 
spathe. The long, pendent terete stems with upturned tips and small leaves 
recall some New Guinea species, in particular R. schlechteri K. Krause and 
I have provisionally included R. cylindrosperma in the Neoguineensis Group 
to which those species are assigned (see Boyce 2000a). 

3. Boyce 345 is provisionally included in R. cylindrosperma although it has 
much more coriaceous leaf laminas with closer venation. It may represent 
an undescribed species but the material is inadequate to make a decision. 

Other specimens seen: SARAWAK, l" Div: Kuching. Hewitt s.n. (SING). BRUNEI 
DARUSSALAM. Belait: Sg. Deriam. Boyce 345 (BRUN, K, L). KALIMANTAN. West 
Kalimantan: Serawai, 8 km NE of Desa Jelundung, Batu Lintang, 1 km S of camp along 
hunting trail. Church, Mahyer & Afriastini 1573 (BO, E, GH, K); Bt. Raya. Nooteboom 
4327 (BO, L); Central Kalimantan: Barito Ulu, P.T. Pamenang logging concession road, 
km 20. Sidyasa PBU 650 (BO, K. L); East Kalimantan: Wanriset research area, road 
Balikpapan to Samarinda, km 15, Sg. Wain area, Ambri & Arifin 354 (BO, L, K). 



Rhaphidophora in Borneo 



35 



4. Rhaphidophora elliptica Ridl. 

Rhaphidophora elliptica Ridl., J. Straits Branch Roy. Asiat. Soc. 44 (1905) 
186; Engl. & K. Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 39—40; 
Merrill, J. Straits Branch Roy. Asiat. Soc, special number (1921) 88. — 
Type: Malaysia, Sarawak, 1 Div., Bau, July 1903, Ridley s.n. (SING, holo). 

Figure 4 

Large, occasionally very large, robust, pachycaul homeophyllous liane to 
30 m; seedling stage a non-skototropic shingling shoot; pre-adult plants 
occasionally forming small terrestrial colonies of appressed shingling shoots; 
adult shoot architecture comprised of greatly elongated, clinging, 
physiognomically unbranched, leafy, non-flowering stems and long, 
moderately elaborated, free, leafy, flowering stems later pendent under 
their own weight; stems smooth, climbing stems broadly rectangular in 
cross-section, the angles often slightly winged, the surfaces between convex, 
free stems spreading, rectangular in cross-section, branching, growing to 
considerable lengths, green, later mid-brown, without fibre at the tips of 
active shoots, internodes to 16 x 1 — 2.5 cm on clinging shoots, usually 
shorter and less stout on free shoots, separated by large oblique leaf scars, 
older stems woody; flagellate foraging stems frequent, often of great length, 
± rectangular in cross-section; clasping roots densely arising from the nodes 
and internodes of clinging stems, densely pubescent; feeding roots rare, 
clinging, pubescent; leaves distichous and ascending on clinging and free 
shoots; cataphylls and prophylls membranous, very soon drying and falling; 
petiole deeply grooved adaxially, 10 — 21.5 x 0.2 — 1 cm, smooth, apical and 
basal genicula weakly defined; petiolar sheath very prominent but soon 
falling, extending to and encircling the apical geniculum, ligulate, very 
soon drying and falling to leave a continuous conspicuous scar from the 
petiole base, around the top of the apical geniculum and back to the base; 
lamina entire, elliptic to elliptic-lanceolate or falcate-oblong, weakly 
channelled along midrib, 16 — 25 x 2.5 — 8 cm, coriaceous, upper surfaces 
semi-glossy, lower surfaces semi-matt, base subacute to rounded or very 
slightly cordate, apex subacute with a somewhat prominent apiculate 
apicule, margins very slightly revolute in dried material; midrib raised 
abaxially, slightly sunken adaxially; primary venation pinnate, slightly raised 
abaxially and adaxially but barely visible in fresh material; interprimaries 
parallel to primaries and only slightly less prominent, very slightly raised 
abaxially and adaxially, ± obscure in fresh material; secondary and tertiary 
venation ± obscure in fresh material, visible as a very faint reticulum in 
dried specimens; inflorescence solitary, subtended by a fully developed 
foliage leaf and one or more membranous, soon falling cataphylls; peduncle 



36 



Gard. Bull. Singapore 53 (2001) 




Figure 4. Rhaphidophora elliptica Ridl. 

A. flowering shoot x : /,, B. leaf lamina x V 3 ; C. venation detail x 3; D. portion of adult sterile 
stem x 7 6 ; E. inflorescence, spathe fallen x l / 2 ; F. spadix detail, post-male anthesis x 4. A - C. E 
- F from Mamit S 37665: D. from Kew LCD acc. no. 1965-49804. 



Rhaphidophora in Borneo 



37 



compressed-cylindric. 4 — 11 x 0.3 — 0.5 cm: spathe canoe-shaped, stoutly 
very short-beaked, 6.5 — 16 x 2 — 3.5 cm, thickly stiff-fleshy, dull yellow, 
paler internally, soon falling at female anthesis; spadix tapering-cylindrical, 
± sessile, inserted level on peduncle. 8 — 14 x 1.5 — 2 cm; stylar region weakly 
rhombohexagonal. 1.8 — 2.2 x 1.9—2.1 mm. ± truncate: stigma punctiform. 
raised, c. 0.3 mm diam.; anthers barely exserted at male anthesis, pollen 
extruded from between ovaries: infructescence 10 — 15 x 2 — 2.5 cm. 

Distribution: Endemic to Borneo. Sarawak (l ' and 3 rd Divisions), Brunei 
Darussalam (Muara. Temburong). Sabah (Sandakan). 

Habitat: On trees and cliffs in primary to disturbed primary lowland 
dipterocarp and swamp forest, on a variety of substrates including limestone 
and shale. Sea level to 115 m altitude. 

Notes: 1. Boyce (1999) treated Rhaphidophora elliptica as a synonym of R. 
montana, and comparison of the description above with that of R. montana 
will reveal rather few morphological differences. However, more detailed 
study of R. elliptica has demonstrated subtle but consistent differences 
between it and R. montana such that I am now recognizing the Bornean 
plants to be a distinct, endemic species. In particular. R. elliptica is distinctive 
in having the leaves distichous and ascending with the lamina shallowly 
channelled along the midrib (leaves spiro-distichous and spreading with 
the lamina flat in R. montana) and in having clinging stems considerably 
wider then deep (c. 4:1 v. c. 2:1 in R. montana). These differences, so 
obvious in the field, are almost completely obscured in herbarium specimens 
unless, as is seldom the case, they are recorded by comprehensive field 
data. 

2. Rhaphidophora elliptica belongs to the taxonomically difficult the 
Hongkongensis Group defined by clinging, occasionally free, stems square 
to rectangular in cross-section, by simple, rather stiff leaves and conspicuous, 
membranous ligulate petiolar sheaths extending to the apical geniculum 
and soon falling to leave a horseshoe-shaped scar at the junction of the 
petiole and lamina. The use of traditional herbarium morphology has proved 
a woefully unsatisfactory method for delimiting species in this group and it 
is hoped that the establishment of an alpha-taxonomy will provide a stimulus 
for further study of the complex using macromolecular data, field 
observations and morphometric techniques. 

Oilier specimens seen: SARAWAK, f Div.: Along road between Kuching and Padawan, 
10 miles SW of main Kuching - Serian highway Croat 53186 (MO); Kuching. Setapok F.R.. 



38 



Card. Bull. Singapore 53 (2001) 



Mamit S 37665 (K, KEP, L, US); One mile west of Bau, Nicolson 1293 (US); Setapok F.R.. 
6 miles south of Kuching. Nicolson 1348 (US); Bau, Purseglove 4473 (K, L, SING); 3 rd Div.: 
Kapit, Upper Batang Rejang, Clemens & Clemens 21929 (GH, K. MO). BRUNEI 
DARUSSALAM. Muara: Hotla 13214 (KYO, L); Temburong: Batu Apoi F.R., ridgetop 
west of Kuala Belalong Field Studies Centre. Poulsen 150 (AAU, BRUN. K). SABAH. 
Sandakan: Sg. Lantoh, ^Lantoh SAN 87848 (K. KEP. L, SAN. SAR); Kinabatangan Road. 
Batu Puteh, Pin-Supu F.R., Bt. Supu, Lim et al. LSP 766 (SAN. SING). 

5. Rhaphidophora elliptifolia Merr. 

Rhaphidophora elliptifolia Merrill, J. Straits Branch Roy. Asiat. Soc. special 
number (1921) 88. — [Rhaphidophora elliptica Ridl., J. Straits Branch 
Roy. Asiat. Soc. 49 (1908) 52, nom. Meg., non Ridl. (1905)]. - - Type: 
Malaysia. Sarawak, l" Div., Kuching, Oct. 3 1905. Hewitt 12 (SING, holo). 

Figure 5 

Large, robust, pachycaul homeophyllous liane to 10 m; seedling stage a 
non-skototropic shingling shoot; pre-adult plants occasionally forming small 
terrestrial colonies of appressed shingling shoots: adult shoot architecture 
comprised of greatly elongated, clinging, physiognomically unbranched, 
leafy, non-flowering stems and long, moderately elaborated, free, densely 
leafy, flowering stems; stems smooth, climbing stems prominently 
rectangular in cross-section, the surfaces slightly convex, free stems 
spreading, irregularly four-angled in cross-section, sometimes irregularly 
terete, with untidy prophyll, cataphyll and petiolar sheath fibre at the tips 
of active shoots, internodes to 7 x 1.5 cm on clinging shoots, shorter and 
less stout on free shoots, separated by large oblique to almost straight leaf 
scars, older stems sub-woody; flagellate foraging stems not observed 
(absent?); clasping roots densely arising from the nodes and internodes of 
clinging stems, pubescent; feeding roots rare, clinging, pubescent; leaves 
spiro-distichous on clinging and free shoots; cataphylls and prophylls 
membranous, soon drying and degrading into netted and tangled fibres; 
petiole deeply channelled adaxially, 10 — 13 x 0.3 — 0.4 cm, smooth, apical 
geniculum well defined, basal genicula rather weakly defined; petiolar sheath 
prominent, extending to and encircling the apical geniculum, briefly ligulate. 
soon drying and degrading into netted fibres, eventually falling to leave a 
continuous slight scar from the petiole base, around the top of the apical 
geniculum and back to the base; lamina entire, narrowly elliptic to narrowly 
elliptic-oblong, 20 — 29 x 4.5 — 7 cm, coriaceous, upper surfaces semi-glossy, 
lower surfaces matt, base acute to decurrent, apex attenuate with a slight 
apiculate apicule, margins revolute in dried material; midrib raised abaxially, 
sunken adaxially; primary venation densely pinnate, raised abaxially and 
adaxially; interprimaries parallel to primaries and barely less prominent, 



Rhaphidophora in Borneo 



39 




Figure 5. Rhaphidophora elliptifolia Men. 

A. portion of adult sterile stem x B. portion of older, leafless stem x 7 3 ; C. leaf lamina x V 2 ; 
D. venation detail x 4:E. inflorescence, spathe fallen x hF.spadix detail, post-female anthesis. 
pre-male anthesis x 8. A & B from Meijer 2I01;C - F from Hewitt s.n. 



40 



Card. Bull. Singapore 53 (2001) 



slightly raised abaxially and adaxially; secondary and tertiary venation ± 
obscure in fresh material, visible as a very faint reticulum in dried specimens: 
inflorescence solitary, subtended by a fully developed foliage leaf and netted 
petiolar sheath, prophyll and cataphyll fibre; peduncle compressed-cylindric, 
5.5 x 0.3 cm; spathe broadly canoe-shaped, stoutly short-beaked, 6 — 6.5 x 
2 — 2.5 cm, thickly stiff-fleshy, dull yellow, persistent into early fruiting; 
spadix cylindrical, ± sessile, inserted level on peduncle, c. 5 x 1 cm; stylar 
region rhombohexagonal, 1.7 — 2 x 1.7 — 2 mm. truncate; stigma rounded, 
raised, c. 0.3 mm diam.; anthers not exserted at male anthesis; infructescence 
c. 7 cm long (known from partial material only). 

Distribution: Endemic to Borneo. Sarawak (I s Division), Kalimantan (East). 

Habitat: Primary dipterocarp forest. Less than 100 m altitude. 

Notes: 1. Rhaphidophora elliptifolia resembles Peninsular Malaysian/ 
Sumateran R. maingayi (see Boyce, 1999) in the persistent fibrous remains 
at the tips of active stems and the stems rectangular in cross-section, but 
differs in the smaller inflorescences (12 — 22 v. 6 — 6.5 cm), the spathe 
persistent into fruiting and the narrower, more coriaceous leaves. 
Rhaphidophora elliptifolia replaces R. maingayi in Borneo. 

2. Confusion with R. elliptica is possible, although the smaller inflorescences 
and persistent fibre at the shoot tips readily distinguish R. elliptifolia. 

Other specimens seen: SARAWAK. No further data, Hewitt s.n. (SING). KALIMANTAN. 
East Kalimantan. Pulau Nunukan, north of Tarakan. Meijer 2101 (L. BO). 

6. Rhaphidophora foraminifera (Engl.) Engl. 

Rhaphidophora foraminifera (Engl.) Engl.. Pflanzenr. 37 (IV.23B) (1908) 
45; Alderw., Bull. Jard. Bot. Buitenzorg III, 4 (1922) 197. — Epipremnum 
foraminiferum Engl, in Bot. Jahrb. Syst. 25 (1898) 11. — Neotype: Malaysia, 
Perak, Taiping, base of Maxwell's Hill Station road, Nicolson 1047 (US, 
neo; L, BH, BO isoneo; designated by Boyce, 1999). 

Figures 6 & 7 

Moderate to large, robust, pachycaul, homeophyllous liane to 15 m; seedling 
stage not observed; pre-adult plants frequently (always?) forming extensive 
terrestrial colonies; adult shoot architecture comprised of elongated, clinging, 
physiognomically unbranched, densely leafy, non-flowering (always?) stems 
and shorter clinging, densely leafy, flowering stems; stems smooth, mid- 




Figure 6. Rhaphidophora foraminifera (Engl.) Engl. 

A. pre-adult climbing shoot x V 4 ; B. pre-adult terrestrial shoot x '/,; C. leaf lamina x '/,. A & B 
from Nicolson 941: C from Boyce 722. 



42 



Gard. Bull. Singapore 53 (2001) 




Figure 7. Rhaphidophora foraminifera (Engl.) Engl. 

A. flowering shoot x B. venation detail x 6; C. inflorescence, spathe fallen x : /,; D. spadix 
detail, male anthesis x 6. A, C & D from Boyce 722; B from Croat 531 18. 



Rhaphidophora in Borneo 



43 



green, with cataphylls and prophylls deliquescing to black mucilage drying 
to leave fragmentary parchment-like remains on petioles, developing 
laminas. inflorescences, internodes 1 — 7 x 0.5 — 3.5 cm. separated by 
prominent straight to slightly oblique leaf scars; flagellate foraging stem 
absent: clasping roots densely arising from the nodes and internodes. 
pubescent; feeding roots clinging, exceedingly robust, densely pubescent 
with dense verticils of prominent corky ramenta: leaves distichous: cataphylls 
and prophylls membranous, soon deliquescing: petiole canaliculate, 22 — 52 
x 0.4 — 1 cm, smooth for the most part but upper 2 — 4 cm and apical 
geniculum densely pubescent, apical geniculum prominent, basal geniculum 
less so; petiolar sheath prominent, extending to apical geniculum, ± short- 
persistent, degrading to weak, slightly netted fibres: lamina entire to slightly 
or extensively perforated, perforations round to rhombic, extending c. + of 
lamina width on each side of the midrib, ovate to oblong-lanceolate or 
oblong-elliptic, slightly oblique, pubescent abaxially when young, 7 — 53 x 
6 — 19 cm. membranous to subcoriaceous, base rounded, acute to slightly 
decurrent, apex acute to acuminate; midrib prominently raised and densely 
pubescent abaxially. ± flush adaxially: primary venation pinnate, slightly 
raised abaxially and adaxially. the leaf appearing slightly quilted, pubescent 
in younger leaves, this indumentum mostly shed in older leaves; 
interprimaries subparallel to primaries, less prominent, slightly raised 
abaxially, slightly impressed adaxially; secondary venation reticulate, slightly 
raised: tertiary venation tessellate. slightly raised: inflorescence mostly 
subtended by one or two large cataphylls. these soon deliquescing into 
black mucilage, this drying and adhering patchily to developing spathe, 
rarely subtended by a ± fully developed foliage leaf; peduncle terete, 3 — 13 
x 0.5 — 1.5 cm: spathe canoe-shaped, stoutly beaked. 10 — 27 x 1.5 — 13 cm, 
stiffly fleshy, dull pale to dark yellow, wide gaping at male anthesis and 
then briefly persistent though maturation of the stamens, eventually falling 
to leave a large ± scar at the base of the spadix: spadix cylindrical, sessile, 
slightly obliquely inserted on peduncle, 7 — 17 x 1.3 — 2.5 cm, dull greenish 
yellow: stylar region moderately developed, rounded to rhombohexagonal, 
1 — 2 x 1 — 1.5 mm. shortly conical when fresh, drying truncate; stigma 
punctiform or elliptic and transversely orientated, raised at male anthesis 
but excavated in dry material, c. 0.3 mm diam.: anthers slightly exserted at 
male anthesis: infructescence 8 — 15 x 2 — 3 cm. dark green before ripening 
to greenish yellow. 

Distribution: Sumatera, Peninsular Malaysia, Borneo - Sarawak (l" and 6 th 
Divisions, but doubtless more widespread), Brunei Darussalam (Belait) 
and Sabah (Pantai Barat). 



44 



Card. Bull. Singapore 53 (2001 ) 



Habitat: On trees, rocks and cliffs in primary to disturbed secondary lowland 
to moist hill dipterocarp forest, on a variety of substrates including limestone 
and shale. 10 — 700 m altitude. 

Notes: 1. Confusion can occur between Rhaphidophora foraminifera and 
R. puberula. They may be distinguished by the position of the inflorescence 
(on short clinging shoots in R. foraminifera, on short free shoots in R. 
puberula), by the presence of black mucilage produced by the deliquescing 
cataphylls and prophylls (present in R. foraminifera, absent in R. puberula), 
in leaf size of mature flowering-sized plants (to 53 x 19 cm in R. foraminifera, 
to 34 x 10.5 cm in R. puberula), by the more or less rounded (R. 
foraminifera) v. oblong (R. puberula) stylar regions, and in overall size of 
the plant (R. foraminifera is a bole climber to 15 m whereas R. puberula 
seldom climbs higher than 3 m and frequently forms large, spreading masses 
on rocks.) 

2. Perforated leaves occur in a number of otherwise rather different-looking 
Rhaphidophora species (e.g., R. foraminifera. R. puberula, R. versteegii) 
suggesting that while a useful diagnostic tool, lamina perforation cannot be 
used to circumscribe taxonomically meaningful groups within 
Rhaphidophora. 

Other Bornean specimens seen: SARAWAK. 1 ' Div.: Road between Kuching and Padawan, 
10 m SW of main Kuching to Serian highway, Croat 53182 (MO); 1 mile from Bau, Nicolson 
1288 (US); 6 th Div: Ulu Balingian, road from Sibu to Bintulu. Boyce 722 (K, M). BRUNEI 
DARUSSALAM. Belait: Labi, Kp Teraja, path along the Sg. Teraja, Boyce 235 (BRUN. 
K, L. MO). SABAH. Pantai Barat: Road from Kota Kinabalu to Tambunan. Ulu Moyog, 
Boyce 1391, 1400 (K, SNP); Tambunan District, Road to Kg Tondulu, Boyce 1414 (K, 
SNP); Moyog District, along road between Kota Kinabalu and Tambunan. 21 miles SE of 
Kota Kinabalu, W slopes of Crocker Range, Croat 53118 (MO); Kinabatangan, Tamegang 
Timber Camp near Kg Pangkaian, Hotta 1416 (KYO, L). 

7. Rhaphidophora korthalsii Schott 

Rhaphidophora korthalsii Schott, Ann. Mus. Bot. Lugd.-Bat. 1(1863) 129; 
Engl, in A. & C. DC, Monogr. Phan. 2 (1879) 246; Engl. & K. Krause in 
Engl., Pflanzenr. 37 (IV.23B) (1908) 49—51, Fig. 21; Alderw., Bull. Jard. 
Bot. Buitenzorg III, 4 (1922) 341. — Type: Indonesia, Java, P.W. Korthals 
s.n. (L, holo; L, P, iso). 

Pothos celatocaulis N.E. Br., Gard. Chron. 13 (1880) 200. — Rhaphidophora 
celatocaulis (N.E. Br.) Alderw., Bull. Jard. Bot. Buitenzorg III, 1 (1920) 
382 & Bull. Jard. Bot. Buitenzorg III, 4 (1922) 198. — Type: Malaysia, 



Rhaphidophora in Borneo 



45 



Sabah. Burbidge s.n.. Hon. Veitch no. 215 (K. holo; K, iso). 

Rhaphidophora maxima Engl.. Bull. Soc. Tosc. Ortic. 4 (1879) 269 & in 
Beccari. Malesia 1 (1882) 271, Tab. xx 1 — 5: Engl. & K. Krause in Engl.. 
Pflanzenr. 37 (IV.23B) (1908) 48^9: Merrill. J. Straits Branch Roy. Asiat. 
Soc, special number (1921) 89. — Type: Sarawak. G. Gading. July 1866. 
Beccari PB 2314 (FI, lecto, selected by Boyce, 1999). 

Rhaphidophora tenuis Engl.. Bot. Jahrb. Syst. 1 (1881) 181 & in Beccari. 
Malesia 1 (1882) 271—272: Engl. & K. Krause in Engl.. Pflanzenr. 37 
(IV.23B) (1908) 53: Merrill. J. Straits Branch Roy. Asiat. Soc. special 
number (1921) 90. — Types: Malaysia. Sarawak. Beccari PB 1977 (FI lecto: 
B isolecto: lecto selected by Boyce. 1999). 

Rhaphidophora korthalsii Schott var. angustiloha Ridl. ex Engl. & K. Krause 
in Engler. Pflanzenr. 37 (IV.23B) (1908) 49: Merrill. J. Straits Branch Roy. 
Asiat. Soc. special number (1921) 89. — Type: Malaysia. Sarawak. Matang. 
July 1903, Ridley s.n. (SING, lecto: lecto: selected by Boyce, 1999). 

Rhaphidophora copelandii Engl., Bot. Jahrb. Syst. 37 (1905) 115: Engl. & 
K. Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 49. — Type: Philippines. 
Mindanao. Davao, Mt Apo. April 1904. Copeland 1193 (PNH. holot; B 
iso). 

Rhaphidophora grandifolia K. Krause. Bot. Jahrb. 44. Beibl. 101 (1910) 11. 
— Type: Philippines, Negros, Negros Orientale. Dumaguete (Cuernos 
Mts). March 1908. Elmer 9464 (PNH, holot; B. E. K. L. LE, MO. iso). 

Rhaphidophora trinervia Elmer, Leafl. Philipp. Bot. 8 (1919) 3073. — Type: 
Philippines. Laguna. Los Banos (Mt Maquiling). June — July 1917. Elmer 
18057 (PNH, holot; FI. K. L. MO. P. iso). 

Rhaphidophora ridleyi Merr.. J. Straits Branch Roy. Asiat. Soc. special 
number (1921) 90. [Rhaphidophora grandis Ridl.. J. Straits Branch Roy. 
Asiat. Soc. 49 (1907) 51, nom. Meg., non Schott 1858 (India) = R. decursiva 
(Roxb.) Schott]. — Tvpe: Malaysia. Sarawak. Tambusan. Sept. 1905. Ridley 
12414 (SING. holo). 

Rhaphidophora latifolia Alderw., Bull. Jard. Bot. Buitenzorg III. 4 (1922) 
341; K. Krause & Alderw.. Nova Guinea 14 (1924) 213. — Type: Indonesia. 
Irian Java. Pionierbivak. 23 July 1920. Lam 711 (BO. holo; L, iso). 



46 



Card. Bull. Singapore 53 (2001) 



Rhaphidophora palawanensis Merr., Philipp. J. Sci. 26 (1925) 451. — Type: 
Philippines, Palawan, Malampaya Bay, Oct. 1922, Merrill BS 11570 (PNH, 
holof ; B, K, P, iso). 

Rhaphidophora trukensis Hosok., J. Jap. Bot. 13 (1937) 195. — Type: 
Federated States of Micronesia, Chuuk (Truk) Island, near Orrip, 29 July 
1939, Hosokawa 8334 (TI , holo). 

[Epipremnum multicephalum Elmer, Leafl. Philipp. Bot. 10 (1938) 3624, 
nom. invai, descr. Angl. — Based on: Philippines, Luzon, Sorsogon, Trosin 
(Mt Bulusan), May 1916, Elmer 16061 (sheets seen at FI. K, L, MO. P. 
PNHt)]. 

Figures 8 & 9 

Very large, occasionally enormous, slender to rather robust, pachycaul. 
heterophyllous liane to 20 m; seedling stage a non-skototropic shingling 
juvenile shoot; pre-adult plants never forming terrestrial colonies; adult 
shoot architecture comprised of greatly elongated, clinging, physiognomically 
unbranched, densely leafy, flowering stems; stems smooth, bright green, 
with sparse to copious prophyll, cataphyll and petiolar sheath fibre, 
especially at the stem tips, internodes to 15 x 3.5 cm, separated by prominent 
oblique leaf scars, older stems subwoody; flagellate foraging stems absent; 
clasping roots densely arising from the nodes and internodes, prominently 
pubescent; feeding roots abundant, clinging and free, very robust, densely 
ramentose-scaly; leaves distichous; cataphylls and prophylls membranous, 
soon drying and degrading to intricately reticulate persistent fibres; petiole 
shallowly grooved, upper part ± terete, (1 — ) 9 — 65 x 0.2 — 1.5 cm, smooth, 
apical and basal genicula prominent; petiolar sheath prominent, 
membranous, strongly to slightly unequal on one side, extending almost to 
or reaching the apical geniculum, of ± short-duration, soon degrading into 
persistent netted fibres, these eventually falling to leave a prominent, slightly 
corky scar; lamina of seedlings overlapping in the manner of roof shingles, 
entire, lanceolate, 5 — 11 x 3.5 — 6 cm, base slightly cordate, lamina of pre- 
adult and adult plants free, entire, pinnatipartite, pinnatisect or pinnatifid, 
10 — 44 x 14 — 94 cm, broadly oblong-elliptic to oblong lanceolate, slightly 
oblique, membranous to chartaceous or subcoriaceous, base truncate and 
very briefly decurrent, apex acute to acuminate, individual pinnae 1 — 10 
cm wide, frequently perforated basally adjacent to the midrib, thus 
appearing stilted; midrib very prominently raised abaxially, slightly sunken 
adaxially; primary venation pinnate, raised abaxially, somewhat impressed 
adaxially, 2 — 4 primary veins per pinna; interprimaries subparallel to 
primaries, slightly raised abaxially, slightly impressed adaxially; secondary 




Figure 8. Rhaphidophora korthalsii Schott 

A. pre-adult shoot x V 4 : B. pre-adult shingling shoot x V 4 . A from Boyce 679; B from Nicolson 
1712. 



48 



Card. Bull. Singapore 53 (2001) 



venation strongly reticulate, slightly raised; tertiary venation invisible; 
inflorescence solitary to several together, first inflorescence subtended by a 
membranous prophyll and one or more cataphylls, these soon degrading 
to netted fibres, subsequent inflorescences subtended by one or more soon 
degrading cataphylls, the whole forming a mass of developing and open 
inflorescences and developing infructescences partially concealed by 
persistent netted cataphyll and prophyll remains; peduncle slightly laterally 
compressed to terete, 6 — 26 x 1—1.5 cm; spathe narrowly canoe-shaped, 
stoutly beaked, 10 — 30 x 3 — 5 cm, stiffly fleshy, greenish to dull yellow, 
gaping wide at female anthesis and then soon falling to leave a large straight 
scar at the base of the spadix; spadix cylindrical, sessile, inserted ± level on 
peduncle, 9 — 26 x 1.5 — 2 cm, dull green to dirty white; stylar region rather 
well developed, mostly rhombohexagonal, 1.5 — 2 x c. 2 mm, slightly conical; 
stigma punctiform to slightly elliptic, if the latter then mostly longitudinally 
orientated, c. 0.3 — 0.5 x 0.2 — 0.4 mm; anthers barely exserted at male 
anthesis; infructescence 14 — 27 x 3 — 3.5 cm, dark green ripening to dull 
orange, stylar tissue abscising to reveal orange ovary cavity pulp. 

Distribution: Widespread in south tropical Asia from Sumatera and southern 
Thailand to Borneo and the Philippines eastwards through the tropical 
western Pacific. In Borneo, widespread in Sarawak, Brunei Darusslam and 
Sabah. In Kalimantan only recorded from East Kalimanatan. 

Habitat: Primary to disturbed secondary lowland, hill and montane forest, 
moss forest, peatswamp forest, on trees, rocks and cliffs on a variety of 
substrates including limestone and ultrabasic soils. 20 — 1800 m altitude. 

Notes: 1. Rhaphidophora korthalsii is a very widespread and variable species, 
hence the extensive synonymy. However, as with Epipremnum pinnatum 
(L.) Engl. (Boyce, 1998) there are several geographical elements that, given 
more intensive study, might warrant formal taxonomic recognition. 
Unfortunately, current herbarium material is inadequate to confirm these 
plants' status and more field observations are needed. 

2. Sterile herbarium material lacking the pre-adult stage may prove difficult 
to distinguish from the Epipremnum pinnatum (in Borneo known wild 
only from east Sabah). Mature leaves of 'typical' E. pinnatum never have 
more than one primary lateral vein per pinna and the stems of R. korthalsii 
lack the prominent irregular whitish longitudinal crests and older stems 
the distinctive matt to sublustrous pale brown papery epidermis typical of 
E. pinnatum. The feeding roots of R. korthalsii are prominently scaly, 
while those of E. pinnatum are lenticellate-corky. The pre-adult stage of R. 



Rhaphidophora in Borneo 



49 




Figure 9. Rhaphidophora korthalsii Schott 

A. flowering shoot, leaves removed x V : ; B. leaf lamina x '/,; C. venation detail x 2; D. 
inflorescence, spathe removed x 1: E. spadix detail, post-male anthesis x 8. All from Kerr 
15051. 



50 



Gard. Bull. Singapore 53 (2001) 



korthalsii is a shingle climber with oblong-elliptic to ovate, slightly falcate 
upwardly pointing leaves overlapping in the manner of roof tiles. 

3. Fertile material of R. korthalsii and Epipremnum pinnatum is readily 
separated by the shape of the style apex (round v. trapezoid) and the 
shape and orientation of the stigma (± punctiform and circumferential v. 
strongly linear and longitudinal) and, if fruits are mature, by seed characters. 
The fruits of R. korthalsii each contain many small ellipsoid seeds with a 
brittle, smooth testa, whereas E. pinnatum has fruits with two large, strongly 
curved seeds with a bony and ornamented testa. 

Other Bornean specimens seen: SARAWAK. 1 Div.: Bau, Brooke 8998 (BM); Lundu, 
Foxworthy 40 (SING); Three miles from Kuching, Haviland & Hose 3605 (K); Kuching. 
Hewitt 40 (SING); Semengoh F.R., 6 miles west of Kuching, Nicolson 1252 (US); 1 mile 
west of Bau, Nicolson 1301 (US): Bako N.P., 20 miles northeast of Kuching, Lintang Path, 
Nicolson 1329 (US); Setapok F.R., 6 miles west of Kuching, Nicolson 1341 (US); Kuching 
Ridlev s.n. (SING); ?> Div.: Hose Mountains, gorge of Sg. Simpurai. Burn & Martin 4926 
(E); Bt. Raya, Kapit, Soepadmo & Chai S28148 (KLU. SAR); 4* Div.: Niah, Ahmed 64 
(SAR. SING): Miri District, Niah, Sg. Sekaloh, G. Subis, Anderson S31956 (E, K, L, SAR, 
SING, US); Kelabit Highlands, Apu Batu Buli, Nooteboom & Chai 2216 (L, SAR, US); 7' h 
Div.: Bintulu, Bt. Urang, Brunig S 12093 (K, SAR): Eastern ridge of Bt. Kans. Bintulu 
district, Hirano & Hotta 1432 (KYO); Sg. Ma'au, Dataran Tinggi Merurong, Tubau, Othman 
et al. S 49050 (K, SAR); 9' h Div.: Tebedu, mile 15. Mohtar el al. S 49245 (K, SAR, US); 
Serian, G. Penrissen; Paie S 16001 (K. LE. SAR). BRUNEI DARUSSALAM. Belait: Ulu 
Ingei, Bt. Batu Patam, lower slopes near Sg. Ingei, Boyce 312 (BRUN, K); Temburong: Sg. 
Temburong above Kuala Belalong, Argent et al. 9136 (E, K); Bt. Biang, Ashton 154 (K, 
US); Sg. Temburong at Wong Nguan gorge, Wong 1727 (BRUN, K). SAB AH. Kudat: 
Ranau, Bt. Kulung, Meijer SAN 122414 (SAN); Ranau. Bt. Kulung. Meijer SAN 122429 
(SAN); Ranau. Kinabalu road to Mesilau, Mikil SAN 38691 (K, SAN); Mesilau, Poore 131 
(K); Sandakan: Paitan F.R., Ampuria SAN 32773 (K, SAN. SING); Elopura, mile 15 Labuk 
Road, Cuadra A 2237 (K, KEP, SAN, SING); Sandakan. Segalid Lokan F.R.. Gibot SAN 
80977 (K, L. SAN); Lamag, Batu Puteh, Majawat & Lasan SAN 88017 (KEP. SAN); Pantai 
Barat: Tambunan, road to Kg Tondulu, Kg Tondulu valley, Boyce 1421 (K, SING, SNP); 
Papar, Kimanis road, Boyce 1443 (K, SNP); Dallas, Clemens & Clemens 26405 (BM, SING); 
Dallas. Clemens & Clemens 26495 (BM, L, UC); Tenompok, Clemens & Clemens 28813 
(BM, K, SING); Upper Kinabalu, Clemens & Clemens 29146 (BM. BO. K. L, SING, UC); 
Upper Kinabalu, Lilau Basin, Mesilau, Meneringtan. ( 'lemens & Clemens 29146a (BM. K. 
L. SING); Penibukan, Clemens & Clemens 31150 (BM, K, SING); Penataran river. Clemens 
& ( lemens 34298 (BM, BO, K, L, UC); Mount Kinabalu N.P.. along road to summit, Croat 
53131 (MO); Lamag. Sg. Pin, Dewol & Harum SAN 89922 (K, SAN); Ulu Segama, Danum 
Valley, path to Rhino Ridge, Dransfield 6302 (K); Pedalaman: Tenom. north ridge of G. 
Malutut, c. 15 km north of Tenom, Kokawa & Hotta 2455 (KYO, L); Tawau: cutting area 
of Luasong Camp, N.B.T. about 60 km NW of Tawau, Kokawa & Hotta 631 (K. KYO), 
Kokawa & Hotta 797 (K. KYO. P); Lahad Datu, Ulu Sg. Sanum, south of camp 111, Stone 
et al. SAN 85269 (KLU, SAN): Lahad Datu, Ulu Sg. Sanum. just north of camp 111. Stone 
et al. SAN 85310 (KLU, SAN). KALIMANTAN. East Kalimantan: Wanariset research 
area, road Samboja to Semoi, km 1 1. Ambri & Arifin 331 (L); Berau, Tandung Redeb, Sg. 
Kelai near Long Lanuk, Kostermans 21125 (BO, L); Berau, near Tandung Redeb, 



Rhaphidophora in Borneo 



51 



Kostermans 21725 (BO. L. US): Danan Pentulak. Bt. Pegah. Main (sub. Polak) 1957 (BO): 
Pulau Nunukan. Tarakan. Meijer 2600 (BO): North of Tarakan. Simengkadu. Meijer 2389 
(BO). 

8. Rhaphidophora latevaginata M. Hotta 

Rhaphidophora latevaginata M. Hotta. Acta Phytotax. Geobot. 22 (1966) 
4. — Type: Malaysia. Sarawak. 4 Division. Bintulu District, about 4 km 
east from Minah Camp. Sg. Kakus. 4 Oct. 1963. Hirano & Hotta 140 (KYO. 
holo). 

Figure 10 

Moderate to very large, robust, pachycaul. homeophyllous neotenic liane 
to 12 m; seedling stage a non-skototropic shingling juvenile shoot: pre-adult 
plants forming small terrestrial colonies of shingling closely appressed leaves: 
adult shoot architecture comprised of clinging, physiognomically unbranched. 
shingling to very densely leafy, sterile stems and almost identical fertile 
stems: stems weakly compressed-terete to weakly rectangular in cross- 
section, smooth, dark green, without prophyll and cataphyll fibre but with 
at least upper parts with thin, black adherent petiolar sheath tissue, 
internodes to 12 x 2 cm. separated by prominent straight scars, but scars 
obscured by leaf bases on all but the oldest stems, lower parts of stem later 
sub-woody with slightly shiny cracking epidermis: flagellate foraging stems 
moderately well developed although often somewhat short and leafy: 
clasping roots arising densely from the nodes and internodes. prominently 
scaly: feeding roots c. 3 mm diam.. brown, minutely pubescent: leaves 
distichous, appressed. ascending and shingling, becoming slightly scattered 
and spreading (often litter-trapping) towards fertile tips: cataphylls and 
prophylls membranous, soon drying black and persisting briefly before 
falling: petiole deeply canaliculate and winged. 3—22 x 0.5 — 2 cm. smooth, 
apical and basal genicula almost obscure in young leaves, later becoming 
prominent, especially the basal geniculum: petiolar sheath very pronounced, 
up to 2.5 cm wide, rather thickly membranous, prominently ligulate. outer 
sheath (away from climbing surface) greatly expanded and partly to 
completely obscuring stem, sheath turning black but persisting some 
considerable time, eventually rotting away to reveal two large scars 
extending to the top of the petiole: lamina broadly oblong-ovate-elliptic. 
8 — 48 x 6.5 — 20.5 cm stiffly coriaceous, bright green, slightly to markedly 
glaucous, base truncate-cordate to broadly cuneate. very briefly decurrent. 
apex rounded to acute with a tiny apicule: midrib prominently raised 
abaxially. slightly sunken adaxially: primary venation pinnate, slightly raised 



52 



(lard. Bull. Singapore 53 (2001) 



abaxially, more so adaxially; interprimaries sub-parallel to primaries, slightly 
raised on both leaf surfaces; secondary venation tessellate-reticulate, slightly 
raised abaxially, ± flush adaxially, all veins much more prominent in dried 
material; inflorescence solitary on a clinging shoot, subtended by a fully 
developed foliage leaf and one or more cataphylls; peduncle laterally 
compressed-cylindrical, 6.5 — 11 x 0.5 — 0.7 cm; spathe not observed; spadix 
stoutly cigar-shaped, sessile; spadix inserted ± level on stipe, 17.5 x 1.5 cm, 
pale green; stylar region rhombohexagonal, c. 2 x 1 mm, truncate; stigma 
slightly raised, elongated, longitudinally orientated, c. 0.75 x 0.2 mm; anthers 
not exserted at male anthesis; infructescence stoutly cigar-shaped, 15 x 2 
cm, stylar region becoming convex at fruit maturity. 

Distribution: Endemic to Borneo. Sarawak (l"' and 7 h Division), Brunei 
Darussalam (Temburong), Sabah (Pantai Barat and Sandakan), Kalimantan 
(localities not traced ). 

Habitat: Primary to secondary moist lowland to hill dipterocarp forest on 
clay and sandstone. 50 — 840 m altitude. 

Notes: 1. Despite its remarkable appearance with densely glaucous-leafy 
shingling shoots ascending, often spirally, high into the canopy, and 
furthermore being widespread and not uncommon in Borneo, R. latevaginata 
was described only relatively recently and, remarkably, is known from 
only 13 herbarium specimens only two of which are more than 50 years 
old. Given the plant's striking appearance, it seems inconceivable that it 
was not noticed by early fieldworkers although quite possibly its overall 
similarity to the juvenile stages of the common and widespread R. korthalsii 
may have led to it being passed by as not worth collecting. 

2. It is not at all clear to what Rhaphidophora latevaginata is related. It is 
the only large neotenic lianescent aroid in Borneo, although neoteny in 
Rhaphidophora occurs in three groups (Boyce, 1999; Boyce & Bogner, 
2000). The similarity of the plant to the juvenile stage of R. korthalsii 
might seem compelling and the shared scaley feeding roots suggest a 
relationship. However, the inflorescences are quite different; the style of 
R. latevaginata is rhombohexagonal and the stigma longitudinally orientated 
(v. style round and stigma ± punctiform in R. korthalsii.) 

Other specimens seen: SARAWAK, f Div.: Matang F.R., 10 miles west of Kuching, Nicolson 
1262 (L, US): 7 Div.: Sibu-Bintulu road. 2.3 km alter bridge over the Batang Kemena. 
Boyce 728 (K, M); Bintulu, about 2 - 4 km east from of Minah Camp. Sg. Kakus. Hirano & 
Hotta 41 (KYO), Hirano & Hotta 149 (KYO); Vicinity of Minah Camp (timber camp of 



Rhaphidophora in Borneo 



53 




Figure 10. Rhaphidophora latevaginata M. Hotta 

A. flowering shoot x : /„; B. adult leaf lamina x '/,; C. venation detail x 3; D. inflorescence, 
spathe fallen x 2 / 3 ; E. spadix detail, early fruiting x 4; F. pre-adult terrestrial shoot x V 3 ; G. pre- 
adult terrestrial shoot x '/,; H. pre-adult terrestrial shoot x '/,. A, D & E from Hirano & Hotta 
140; B & C from Dewol & Meijer 131653; F & G. from Hirano & Hotta 149; H from Hirano & 
Hotta 41. 



54 



Gard Bull. Singapore 53 (2001) 



L.L.B.T.C.), Sg. Kakus. Hotta 15717 (KYO). BRUNEI DARUSSALAM. Temburong: Sg. 
Temburong at Kuala Belalong, Boyce 402 (BRUN, K, L, SING, MO); Batu Apoi F.R.. 
ridge west of Kuala Belalong Field Studies Center slope, between Danish Plot and Sg. 
Belalong, Poulsen 165 (AAU, BRUN, K). SAB AH. Pantai Barat: Road from Kota Kinabalu 
to Tambunan, Ulu Moyog, Boyce 1389 (K, SNP); Sandakan: Phenological trail above sawmill, 
Dewol & Meijer SAN 131653 (SAN); Kinabatangan District, Kg. Bilit, Bt. Bilit. Lim el al. 
1272 (SING). KALIMANTAN. Locality not traced: Batu Makele. Batu Eilander, Raap 118 
(BO); G. Dansu, Pentaclale, Pegah, Main (sub. Polak) 1958 (BO). 

9. Rhaphidophora lobbii Schott 

Rhaphidophora lobbii Schott, [Bonplandia 5(2) (1857) 45, nom. mid.], Prodr. 
Syst. Aroid. (1860) 379; Engl, in A. & C. DC, Monogr. Phan. 2 (1879) 
240; Engl. & K. Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 33—34; 
Merrill, J. Straits Branch Roy. Asiat. Soc. special number (1921) 89. — 
Scindapsus lobbii (Schott) Ender, Index Aroid. (1864) 74. — Type: 
Singapore, Lobb 44 (K, holo). 

Figure 11 

Small to moderate, somewhat slender, leptocaul, homeophyllous liane to 5 
m; seedling leafy at germination and skototropic by alternating series of 
congested leafy and elongated leafless shoots; pre-adult plants forming 
diffuse terrestrial colonies; adult shoot architecture comprised of greatly 
elongated, clinging, physiognomically unbranched, leafy, non-flowering 
stems and long moderately elaborated, free, densely leafy, flowering stems; 
stems puberulent-scabrid to asperous, especially on older growth, climbing 
stems weakly rectangular to ± terete in cross-section, free stems ± terete in 
cross-section, often branching extensively and growing to moderate lengths 
pendent under their own weight, dull brown, without prophyll, cataphyll 
and petiolar sheath fibre, internodes to 13 x 0.6 cm on clinging and free 
shoots, flowering shoots with much shorter internodes, separated by weak 
to rather prominent, slightly oblique leaf scars, older stems woody; flagellate 
foraging stems absent; clasping roots sparsely arising from the nodes and 
internodes of clinging stems, densely pubescent; feeding roots rather rare, 
sometimes clinging but often free, pubescent; leaves weakly spiralled and 
often sparsely arranged on clinging and proximal portions of free shoots, 
densely spirally-distichous distally on flowering shoots; cataphylls and 
prophylls membranous, very soon drying and falling; petiole grooved 
adaxially, 4 — 9.5 x 0.2 — 0.3 cm, smooth, with a moderate apical and 
prominent basal geniculum; petiolar sheath slightly prominent, extending 
beyond the apical geniculum by two ligules, very soon drying and falling in 
strips to leave a continuous scar from the petiole base, around the top of 
the apical geniculum and back to the base; lamina entire, narrowly elliptic 



Rhaphidophora in Borneo 



5? 




Figure IL Rhaphidophora lobbii Schott 

A. portion of stem with two flowering shoots x .: B. pre-adult terrestrial shoots x 7 3 ; C. stem 
detail x 3: D. leaf lamina x -7 4 : E. venation detail x 4: F. inflorescence, spathe fallen x 2: G. 
spadix detail, early fruiting x 8. A & C from Kunstler ( Dr King's Collector') 10571: B from 
Boyce 995: D & E from Nauen s.n.: F & G. from Alvins 270. 



56 



Card. Bull. Singapore 53 (2001 ) 



to elliptic-lanceolate to oblong or oblanceolate, 6 — 24 x 2 — 10 cm, very 
softly coriaceous, upper surfaces slightly glossy, lower surfaces pale satin- 
matt, drying markedly discolorous, dark brown above, pale brown below, 
base cuneate to acute or subovate, briefly decurrent, apex acute to ovate- 
acuminate, with a prominent apiculate apicule; midrib slightly raised 
abaxially, slightly sunken adaxially; primary venation pinnate, slightly raised 
abaxially, prominent (dark veins against pale lamina) in dried material; 
interprimaries parallel to, but much less distinctive than, primaries, very 
slightly raised abaxially; secondary and tertiary venation ± invisible in fresh 
material, barely visible in dried specimens, reticulate; inflorescence solitary, 
subtended by a fully developed foliage leaf and a very soon falling cataphyll; 
peduncle compressed-cylindric. 1.5 — 5 x 0.15 — 0.4 cm; spathe cigar-shaped, 
stoutly long-beaked, 3 — 5 x 0.4 — 1 cm, thickly fleshy, exterior minutely 
puberulent, dull green to yellowish, soon falling at female anthesis to leave 
a substantial, slightly oblique, scar; spadix slender cylindrical, sessile, inserted 
level on peduncle, 3 — 3.5 x 0.4 — 0.5 cm, dull yellow-white; stylar region 
rather well developed, mostly rhombohexagonal, 1.9 — 2.4 x c. 2 mm, 
truncate; stigma punctiform, c. 0.3 mm diam., prominent in dried material; 
anthers barely exserted at male anthesis, pollen extruded from between 
ovaries; infructescence oblong-cylindric, 2.5 — 4 x 1 — 1.2 cm. 

Distribution: Southern Thailand, Peninsular Malaysia, Singapore, Sumatera, 
North Borneo and into Sulawesi. 

Habitat: Primary to disturbed secondary lowland dipterocarp and 
peatswamp forest on wet to inundated soils. Sea level to 250 m altitude. 

Notes: 1. A climber distinctive by the slender, asperous stems, softly leathery 
leaves and minutely puberulent spathe exterior, a combination of characters 
unknown in any other Malesian Rhaphidophora. In dry material the strongly 
discolorous leaves, combined with the primary veins prominently darker 
than the abaxial leaf surface, are unmistakable. 

2. In Borneo, confusion with R. cylindrosperma is possible although the 
latter differs in its more coriaceous leaves, different leaf venation and in 
having a longer, externally glabrous spathe. 

3. Rhaphidophora lobbii is habitually found in wet to inundated ground, 
an unusual habitat for a monsteroid liane, cf. Epipremnum amplissimum 
Schott, and E. giganteum (Roxb.) Schott. 

Other Bornean specimens seen: SARAWAK, l" Div.: Sctapok F.R., 4 miles south of Kuching. 



Rhaphidophora in Borneo 



57 



Anderson S 23496 (SAR): Semengoh F.R.. 6 miles west of Kuching. Nicolson 1258 (US): 
Setapok F.R.. 6 miles south of Kuching. Sicolson 1359 (US); 2 Div.: Simanggang. Triso 
P.F.. Anderson 14547 (SING ): 8 th Div.: Binatang. Pulau Bruit. Anderson 8032 (SING): Sg. 
Kelepu. Pulau Bruit. Anderson 8007 (L. SING): Baram. Anderson S 3084 (SAR. SING). 
BRUNEI DARUSSALAM. Temburong: Selapon. village margin. Dransfield 6917 (BRUN. 
K. L): Batu Apoi F.R.. on ridge between Kuala Belalong and Bt. Belalong. in permanent 
Plot 2. Poulsen 362 (AAU. BRUN. K): Belait: Melilas, Ulu Ingei. path from Ulu Ingei to 
Sg. Tupai. Sands 5919 (BRUN. K. L): Rasau. Van Niel 4250 (L. MO). SABAH. Sandakan: 
Sepilok F.R.. Gambating SAS 94588 (K. L. SAN): Sg. Dagat. George et al. SAN 120521 (K. 
SAN): Kinabatangan. Tamegang Timber Camp near Kg. Pangkaian. Kokawa & Horta 1417 
(KYO. L): Labuk Road. Meijer SAN 38784 (K. L. SAN): Below waterfall Kebun Cina F.R.. 
Meijer & Dewol SAN 131749 (SAN): Bongaya F.R.. Kodoh & Aban SAN 82024 (K. L. 
SAN. SAR. SING). KALIMANTAN. West Kalimantan: Pontianak. Polak 308 (BO): Sg. 
Banghong. Polak 645 (BO. K): Bt. Raya. Tumbang Riang. Veldkamp 7910 (BO. L). East 
Kalimantan: Wanariset. Balikpapan. Afriastini 116 (BO): Km 79 on road from Sangui Sg. 
Mantaya. Kab. Kotawaringin Timur. Argent et al. 93138 (B. E ): Berau. Tandung Redeb. Sg. 
Kelai. near Long Lanuk. Kostennans 21133 (BO. L): Munukan. north of Tarakan. Meijer 
2034 (BO): Borneo*. Korthals 149 (L) & Korthals s.n. (L). 

10. Rhaphidophora megasperma Engl. 

Rhaphidophora megasperma Engl.. Bot. Jahrb. Syst. 25 (1881) 8: Engl. & 
K. Krause in Engl.. Pflanzenr. 37 (IV.23B) (1908) 29. Fig. 9: Aldenv.. Bull. 
Jard. Bot. Buitenzorg III. 1 (1920) 383: Merrill. J. Straits Branch Roy. 
Asiat. Soc. special number (1921) 89. — Type: Indonesia. Kalimantan. 
Central Kalimantan. Tumbang Hiang. 2 Sept. 1881. Grabowski 48 (B. holo). 

Scindapsus havilandii Ridl.. J. Straits Branch Roy. Asiat. Soc. 44 (1905) 
184: Engl. & K. Krause in Engl.. Pflanzenr. 37 (IV23B) (1908) 75: Merrill. 
J. Straits Branch Roy. Asiat. Soc. special number (1921) 91. synon. no*. — 
Type: Malaysia. Sarawak, 2 nd Div.. Saribas. July 1892. HavUand 2089 (K. 
holo: SING. iso). 

Rhaphidophora jaculiformis Aldenv.. Bull. Jard. Bot. Buitenzorg III. 4 
(1922) 197. synon. nov. — Type: Indonesia. Kalimantan, no further data. 
1893. Jaheri s.n. (BO. holo: BO. iso). 

Rhaphidophora subfalcata M. Hotta. Acta Phytotax. Geobot. 22 (1966) 6. 
synon. nov. — Type: Malaysia. Sarawak. 7 Div.. Bintulu. along Sg. Kakus 
from Sg. Tmggili to Sg. Puteh. 13 Nov. 1963. Hirano & Hotta 1068 (KYO. 
holo). 

Figure 12 

Moderate sized, slender to slightly robust semi-leptocaul homeophyllous 
creeping and climbing liane to 5 m: seedling stage not observed: pre-adult 



58 



Gard Bull. Singapore53 (2001) 



plants forming extensive terrestrial colonies; adult shoot architecture 
comprised of greatly elongated, clinging, physiognomically unbranched, 
sparsely leafy, non-flowering stems and short, little-branched, free, densely 
leafy, flowering stems; stems smooth, terete in cross-section, internodes to 
10 x 1.7 cm on clinging shoots, much shorter on free shoots, separated by 
slightly oblique leaf scars, older stems subwoody; flagellate foraging stems 
absent; clasping roots sparsely arising from the nodes and internodes of 
clinging stems, pubescent; feeding roots rare, clinging, pubescent; leaves 
spiro-distichous on clinging shoots, distichous on free shoots; cataphylls 
and prophylls membranous, soon drying chartaceous and persisting at the 
tips of flowering shoots, then falling; petiole grooved canaliculate, 1 1 — 29 x 
0.2 — 0.4 cm, apical and basal geniculum moderately prominent; petiolar 
sheath wide and membranous but very soon falling and thus usually not 
prominent, extending almost to or reaching apical geniculum, very soon 
drying and degrading into very sparse, soon-falling fibres; lamina entire to 
slightly perforated, perforations round to rhombic, extending c. V 4 — V 2 of 
lamina width on each side of the midrib, oblong-lanceolate or oblong- 
elliptic, oblique, 12.5 — 42 x 3 — 10.5 cm, thinly coriaceous, base unequal, 
rounded, weakly acute to subrounded, apex acuminate; midrib prominently 
raised abaxially, ± sunken adaxially; primary venation pinnate, slightly raised 
abaxially and adaxially; interprimaries subparallel to primaries, much less 
prominent, slightly raised abaxially and adaxially; secondary venation 
prominently reticulate, raised abaxially, less so adaxially; inflorescence two, 
three or more, together, each subtended by a prominent chartaceous 
prophyll and one or more chartaceous cataphylls; peduncle slender to 
somewhat stout, terete, 4 — 1 x 0.2 — 0.3 cm; spathe broadly canoe-shaped, 
stout-beaked, 4 — 6 x 2 — 3.5 cm, stiff-fleshy, dull yellow on opening, 
persistent into early fruiting, eventually falling to leave a prominent scar; 
spadix stoutly to somewhat slender cylindrical, sessile, inserted obliquely 
on peduncle, 2.5 — 4 x 0.5 — 0.7 cm, dull cream; stylar region mostly 
hexagonal, 1.1 — 1.2 x 1 — 1.1 mm; stigma punctiform, very prominent, 0.1 — 
0.2 x c. 0.3 mm; anthers exserted at male anthesis; infructescence stoutly 
oblong-cylindrical, 3.5—5 x 1.2 — 1.5 cm. 

Distribution: Endemic to Borneo (Sarawak (\ \ 3 rd , 7"' and 8 h Divisions) 
and Kalimantan (West and Central). 

Habitat: Primary to disturbed secondary lowland, often riverine, forest on 
clay-loam. 40 — 130 m altitude. 

Notes: 1. Distinctive in Borneo by the short leafy free side shoots bearing 
clusters of inflorescences subtended and interspersed by prominent 



Rhaphidophora in Borneo 



59 




Figure 12. Rhaphidophora megasperma Engl. 

A. flowering shoot x V,; B. leaf lamina x '/,; C. venation detail x 2; D. leaf lamina x 7 3 ; 
inflorescence, spathe removed x 1V 2 ; E. spadix detail, post-male anthesis x 6: spadix detail, 
early fruiting x 4. A & F from Ridley 14444; B— E from Burkill 6267. 



60 



Card. Bull. Singapore 53 (20(H) 



chartaceous prophylls and cataphylls and by the spathe drying and persistent 
into early fruiting. The clustered inflorescences subtended by chartaceous 
cataphylls recall some New Guinea species, notably Rhaphidophora 
versteegii. 

2. Boyce (1999) merged R. megasperma with R. puberula. However, study 
of further Bornean specimens of both taxa clearly shows that this was in 
error and that R. megasperma is the earliest name for a Bornean endemic 
hitherto called Scindapsus havilandii, Rhaphidophora jaculiformis and R. 
subfalcata. 

3. Hotta's name R. subfalcata has been applied to large plants of R. 
megasperma with perforated leaves. The presence of perforated and 
unperforated leaves in individuals of same species is not uncommon in 
Rhaphidophora; R. puberula is typical of such a phenomenon. 

4. Both type sheets of Ridley's Scindspaus havilandii are fertile; so it is odd 
that Ridley should have considered it to belong to Scindapsus since the 
generic diagnostic characters, number and position of ovules (solitary and 
basal v. many on one or two parietal placentas), are readily observable. 

5. The types of Aldrwerelt's R. jaculiformis match R. megasperma perfectly. 

Other specimens seen: SARAWAK. 1 Div.: Matang. Ridley s.n. (K); 3 Div.: Kapit. Sung, 
upper Sg. Katibas. Sg. Joh trail. Zainudin et al. 4537 (K, UKMB); 7 Div.: Sg. Bejangung. a 
branch of the Sg. Anap, Bintulu District, Hotta 15578 (KYO); 8' h Div.: Bandar Sri Aman - 
Sibu road, 156 km before Sibu below bridge over the Sg. Undup. Boyce 721 (K. M); Serian. 
Kg. Tebekang, Othman & Munting S55097 (K, L, SAR). KALIMANTAN. West Kalimantan: 
Serawai, 8 km NE of Desa Jelundung, Batu Lintang, next to camp. Church. Mahyer & 
Afriastini 1668 (BO, GH, K); Serawai, Winkler 238 (E, L); Serawai. near Talok Nibung. 
Winkler 1250 (E, L). Lake Tajan. Main (sub. Polak) 1758 (BO, L); Upper Samba river, 60- 
80 km NNW of Tumbang Samba, Tumbang Habangoi to Tumbang Riang, Mogea 3550 
(BO, K, KEP. L); Central Kalimantan: Bt. Raya, Nooteboom 4120 (L, BO); 

11. Rhaphidophora minor Hook.f. 

Rhaphidophora minor Hook./., Fl. Brit. Ind. 6 (1893) 544; Engl. & K. 
Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 21, Fig. 4; Merrill, J. Straits 
Branch Roy. Asiat. Soc, special number (1921) 89. — Type: Malaysia, 
Malacca, Griffith 5988 (K, lecto; K, P, isolecto, selected by Boyce, 1999). 

Rhaphidophora celebica K. Krause, Notizbl. Bot. Gart. Berlin-Dahlem. 11 
(1932) 331. — Type: Indonesia, Sulawesi, northwest of the island, near 



Rlwphidophora in Borneo 



61 



Toli-Toli, Jan. 1910. Schlechter 20698 (B. holo). 

[Rhaphidophora palawanensis Furtado. nom. mid. in sched. var., non. Merr. 
(1925)] 

Figure 13 

Small to moderate, slender, leptocaul. homeophyllous liane to 6 m: seedling 
not observed; pre-adult plants forming diffuse terrestrial colonies; adult 
shoot architecture comprised of greatly elongated, clinging, physiognomically 
unbranched. leafy, non-flowering stems and long moderately elaborated, 
free, densely leafy, flowering stems: stems smooth, flexuous. climbing stems 
± terete, occasionally weakly 4-angled in cross-section, free stems somewhat 
laterally compressed in cross-section, often branching extensively, growing 
to considerable lengths and pendent under their own weight with flowering 
tips upturned, without prophyll, cataphyll and petiolar sheath fibre, 
internodes to 13 x 1.2 cm on clinging and free shoots, flowering shoots with 
much shorter internodes, separated by weak straight leaf scars, older stems 
woody: flagellate foraging stems absent; clasping roots arising sparsely or 
singly from the clinging stems, pubescent: feeding roots solitary from nodes, 
free, stout, slightly pubescent; leaves weakly distichous and sparsely arranged 
on clinging and proximal portions of free shoots, moderately densely 
distichous distally on flowering shoots; cataphylls and prophylls 
membranous, very soon drying and falling: petiole grooved adaxially, 3 — 6 
x 0.1 — 0.25 cm, smooth, with a slight apical and prominent basal geniculum: 
petiolar sheath slightly prominent, extending beyond the apical geniculum 
by two small ligules, very soon drying and falling in strips to leave a 
continuous scar from the petiole base, around the top of the apical 
geniculum and back to the base; lamina entire, narrowly falcate-elliptic to 
falcate-lanceolate or falcate-oblanceolate. 2.5 — 16 x 1.2 — 3 cm. thinly 
coriaceous, drying pale straw-coloured, base cuneate to acute or subovate, 
apex acute with a prominent apicule: midrib raised abaxially. slightly raised 
adaxially; primary venation pinnate, slightly raised on both surfaces 
prominent (raised) in dried material: interprimaries subparallel to, but much 
less distinctive than, primaries, sometimes degrading into weakly reticulate 
venation, very slightly raised abaxially: secondary and tertiary venation ± 
invisible in fresh material, barely visible in dried specimens, reticulate; 
inflorescence solitary, subtended by a fully developed foliage leaf and a 
very soon falling cataphyll: peduncle compressed-cyhndric, 3 — 4 x 0.3 — 0.5 
cm: spathe cigar-shaped, stoutly long-beaked, 3 — 9 x 1 — 1.5 cm. thin, dull 
green to dull yellow, soon falling at female anthesis to leave a large, straight, 
scar; spadix slender cylindrical, sessile, inserted level on peduncle. 2.5 — 7 x 
0.5 — 0.6 cm. dull yellow-white; stylar region rather well developed, mostly 



62 



Gard Bull. Singapore 53 (2091) 



rhombohexagonal, 1.4 — 2 x c. 2 mm, truncate; stigma punctiform, c. 0.3 
mm diam., slightly prominent in dried material; anthers well-exserted at 
male anthesis; infructescence oblong-cylindric. 4.5 — 7 x 1 — 2.5 cm. 

Distribution: Southern Thailand. Peninsular Malaysia, Singapore, Sumatera, 
throughout Borneo (widely scattered), Sulawesi and into the Philippines 
(Mindanao, Palawan). 

Habitat: Lowland peatswamp forest, riverine forest, rear of freshwater 
mangrove. Sea level to 10 m altitude. 

Notes: 1. In the fresh state confusion with Rhaphidophora sylvestris is 
possible, although the thinner, more prominently veined leaf and longer 
spathe beak of R. minor Hook./, are diagnostic. Dried material of R. minor 
is notable for the uniformly pale straw coloured leaves. 

2. In Borneo, R. minor is habitually found in swampy habitats whereas 
elsewhere in its range it tends to be in drier places. 

Other Bornean specimens seen: BRUNEI DARUSSALAM. Belait: Kuala Belait. Van Niel 
3944 (L); Seria, Van Niel 4138 (L); Sg. Belait. Van Niel 4634 (L). SARAWAK, l" Div.: 
Kuching. Hewitt s.n. (K): Kuching, Ridley s.n. (SING); Kuching, Ridley s.n. (K). 7' h Div.: 
Ulu Bawan, Balingian, Ashton S 19585 (GH, K. L, MO, SAR, SING, US); Sg. Tengah, 
Brooks 1016 (BM); Batang Lassa, near Rhuma Jumbau. Mamit S 33623 (K, L. MO. SAR. 
US). SAB AH. Sandakan: Beluran, Sg. Mangkayok, near Sg. Kalagan, Meijer SAN 51617 
(SAN); Ulu Dusum, Meijer SAN 122736 (E, SAN, US). KALIMANTAN. West Kalimantan: 
Mempaura, Teysmann 8105 (BO); Central Kalimantan: Sampit, Buwalda 7917 (BO); East 
Kalimantan, 20 km north of Wanariset, Kg. Padang, Samboja, Sidiyasa 1111 (K, L); Cape 
Genderen. Pladjoe, Amdjah 7 (BO. K. L. SING), Amdjah 32 (BO, L), Amdjah 951 (BO, K, 
L); G. Samenggaris, Amdjah 1084 (BO, K, L): South Kalimantan: no further data, Winkler 
3448 (L); Locality not traced: Hallier35, 66 (BO); Long Lansiah (?), Teysmann 11338 (BO, L); 

12. Rhaphidophora puberula Engl. 

Rhaphidophora puberula Engl., Bot. Jahrb. Syst. 1 (1881) 180 & in Beccari, 
Malesia 1 (1882) 269—270; Engl. & K. Krause in Engl., Pflanzenr. 37 
(IV.23B) (1908) 30. — Type: Indonesia, Sumatera, West Sumatera, Padang, 
Ajer Mantjoer, Aug. 1878, Beccari s.n. (FI, holo; B, iso). 

Rhaphidophora scortechinii Hook./., Fl. Brit. India 6 (1893) 545; Engl. & 
K. Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 23. — Type: Malaysia, 
Perak, Scortechini 347 (K, holo; CAL, iso). 

Rhaphidophora kunstleri Hook./., Fl. Brit. Ind. 6 (1893) 546; 548; Engl. & 



Rhaphidophora in Borneo 



63 




Figure 13. Rhaphidophora minor Hook/. 

A. portion of adult, sterile shoot x '/,; B. flowering shoot x l / 2 ; C. leaf lamina x 2 / 3 ; D. venation 
detail x 2; E. inflorescence, spathe just beginning to abscise x 1 1 / 2 ; F. spadix detail, post-female 
anthesis, pre-male anthesis x 8; G. spadix detail, early fruiting x 5. A & D from Winkler 238; B 
& from Church et al. 1668; E & F from Hotta 1068. 



64 



Card. Bull. Singapore 53 (2(101) 



K. Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 30; Alderw., Bull. Jard. 
Bot. Buitenzorg III, 4 (1922) 338. — Type: Malaysia, Perak, Bt. Larut, July 
1883, Kunstler 4538 (K, lecto, selected by Boyce, 1999). 

Rhaphidophora gracilipes Hook./., Fl. Brit. Ind. 6 (1893) 545; Engl. & K. 
Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 30, Fig. 10; Alderw., Bull. 
Jard. Bot. Buitenzorg III, 1 (1920) 384. — Type: Malaysia, Perak, June 
1886, Kunstler 10271 (K, lecto; BM, SING, isolecto, selected by Boyce, 
1999). 

Rhaphidophora batoensis Engl. & K. Krause in Engl., Pflanzenr. 37 (IV.23B) 
(1908) 27. — Type: Indonesia, Sumatera, North Sumatera, Kepulauan Batu, 
25 Sept. 1896, Raap 370, (B, lecto; BO, isolecto, selected by Boyce, 1999). 

Rhaphidophora hallieri Alderw., Bull. Jard. Bot. Buitenzorg III, 1 (1920) 
385; Alderw., Bull. Jard. Bot. Buitenzorg III, 4 (1922) 196. — Type: 
Kalimantan, Hallier 1187 (BO, holo; BO iso). 

Rhaphidophora pilosula Alderw., Bull. Jard. Bot. Buitenzorg III, 1 (1920) 
386; Alderw., Bull. Jard. Bot. Buitenzorg III, 4 (1922) 196. — Type: Cult. 
Bogor Bot. Gard, June 1919, Alderwerelt s.n. (BO, holo). 

Figure 14 

Moderate to large, rather robust, semipachycaul homeophyllous liane to 5 
m; seedling stage not observed; pre-adult plants often forming small 
terrestrial colonies; adult shoot architecture comprised of elongated, clinging, 
physiognomically unbranched, leafy, non-flowering stems and short, usually 
unbranched, free, densely leafy, flowering stems; stems smooth, terete in 
cross-section, with very sparse prophyll, cataphyll and petiolar sheath fibre, 
this soon falling, internodes to 13 x 1.5 cm on clinging shoots, usually 
shorter and stouter on free shoots, separated by large, straight, corky leaf 
scars, older stems woody; flagellate foraging stems absent; clasping roots 
sparsely arising from the nodes and internodes of clinging stems, pubescent; 
feeding roots rare, clinging, pubescent; leaves distichous on clinging and 
free shoots; cataphylls and prophylls membranous, soon drying and 
degrading into very sparse fibres, these soon falling; petiole deeply grooved 
to basally canaliculate, 9 — 33 x 0.2 — 3 cm, with the distal portion and 
apical geniculum minutely pubescent, especially when newly expanded, 
apical and basal geniculum moderately prominent; petiolar sheath very 
prominent, extending to just below the apical geniculum, soon drying and 
degrading into sparse, soon-falling fibres; lamina entire to slightly 
perforated, perforations round to rhombic, extending c. % — 7 4 of lamina 



Rhaphidophora in Borneo 



65 



width on each side of the midrib, ovate to oblong-lanceolate or oblong- 
elliptic, oblique to falcate, minutely pubescent abaxially when young, 8 — 
34 x 3.5 — 10.5 cm, thinly coriaceous to subchartaceous, base unequal, 
rounded, acute to slightly decurrent, apex acute to acuminate; midrib 
prominently raised and pubescent abaxially. + sunken adaxially; primary 
venation pinnate, slightly raised abaxially and adaxially, the leaf appearing 
slightly quilted, pubescent in younger leaves, indumentum mostly shed in 
older leaves; interprimaries subparallel to primaries, very slightly less 
prominent, slightly raised abaxially, slightly impressed adaxially; secondary 
venation tessellate to weakly reticulate, slightly raised; tertiary venation not 
visible; inflorescence solitary, very rarely two together, fragrant of frangipani 
and pineapple {fide Hay et al. 9029) subtended by a fully developed foliage 
leaf and, if more than one inflorescence, then second preceded by a large 
cataphyll: peduncle strongly compressed-cylindric, 4 — 9 x 0.4 — 0.6 cm; spathe 
canoe-shaped, stoutly long-beaked, 5.5 — 19 x 1 — 6.5 cm. stiff-fleshy, dull 
green to dull yellow, pale yellow to apricot-orange internally, falling (after?) 
female anthesis to leave a large, straight scar; spadix very weakly clavate 
cylindrical, sometimes slightly curved, sessile, inserted + level on peduncle, 
3 — 11.5 x 1 — 1.6 cm, cream; stylar region mostly hexagonal, 1.5 — 2 x 2.1 — 
2.5 mm, truncate; stigma punctiform, prominent, 0.5 — 0.75 x 0.45 — 0.75 
mm; anthers exserted at male anthesis; infructescence stoutly oblong- 
cylindrical. 4.5 — 14 x 1.5 — 2 cm, glaucous. 

Distribution: Sumatera, Peninsular Malaysia. Nusa Tenggara and throughout 
Borneo. 

Habitat: On trees and rocks in primary and secondary lowland to lower/ 
upper montane transition forest, often on steep slopes, on granite and 
limestone. 90 — 1550 m altitude. 

Notes: 1. Given the overall uniformity of this species, it is quite extraordinary 
that Rhaphidophora puberula should have been redescribed no fewer than 
seven times based mostly on minor differences of leaf shape. 

2. Superficially similar to R. foraminifera, especially in the perforate-leaf 
forms, R. puberula can readily be distinguished from R. foraminifera by 
the production of inflorescences on free lateral shoots and in having the 
active shoot tips without the characteristic black mucilage of R. foraminifera. 
Generally, flowering plants of the perforate-leaved forms of R. puberula 
have smaller leaves that are noticeably less perforated than those of R. 
foraminifera. 



66 



Card. Bull. Singapore 53 (2001) 



Other Bornean specimens seen: SARAWAK. 1 ' Div.: Matang, Ridley s.n. (K); 4* Div.: 
Baram District, G. Mulu N.P., Hidden Valley, Argent et al. 887 (L); i Div.: Bintulu, Segan 
F.R., Chai S31999 (K, L, SAN, SAR, SING, US). BRUNEI DARUSSALAM. Belait: Labi, 
track to Rampayoh waterfall, Ahmad 15 (BRUN, K). SABAH. Sandakan: Bettohan near 
Sandakan, Kloss SFN 19099 (K, L, SING); Lamag, Kinabatangan, Ulu Sg. Pin, Madani & 
Saigol SAN 90180 (K, L, SAN, SAR); Pantai Barat: Keningau, Sg. Labou, Mantor SAN 
110980 (GH, K, SAN); Tuaran District, below summit of Lawa Mandau, Telekom station 
east of Telipok, Meijer et al. SAN 136186 (GH, SAN); Pedalaman: Kiau. Mount Kinabalu, 
Clemens 10142 (UC); Gurulau Spur, Mount Kinabalu Clemens 10772 (UC); Dallas, Clemens 
& Clemens 26453 (BM. K, L. SING); Dallas, Clemens & Clemens 26781 (BM); Dallas, 
Clemens & Clemens 26876 (BM, BO, GH, K, L, SING, UC); Tenompok, Clemens & 
Clemens 26900 (BM, BO, K, L, SING, UC); Kinabalu N.P., along the road between Park 
H.Q. and Tenompok, Kokawa & Hotta 3089 (KYO); Tawau: Elmer 21593 (BO, K, L, MO, 
SING, UC); Sabah Foundation logging area Umas-Umas, Krispinus SAN 87213 (K, KEP, 
L, SAN, SAR); Kalabakan, km 41 Imbak road, Luasong, Krispinus SAN 95680 (K, L, SAN, 
SAR); Lahad Datu, Madai Caves, Madai Baturong F.R., Kokawa & Hotta 1097 (KYO, L). 
Locality not traced: Pandawan, Sg. Mesopo, Fidilis & Sumbing SAN 113389 (K, KEP, 
SAN); KALIMANTAN. Cult. Bogor, Y70, from Kalimantan collection, Ashton 12626 (BM, 
BO) & Nicolson 942 (BO, L, US). 

13. Rhaphidophora sylvestris (Blume) Engl. 

Rhaphidophora sylvestris (Blume) Engl, in A. & C, DC, Monogr. Phan. 2 
(1879) 239 & in Beccari, Malesia 1 (1882) 268; Engl. & K. Krause in Engl., 
Pflanzenr. 37 (IV.23B) (1908) 22—25, Fig. 6 {'silvestrisj, Alderw., Bull. 
Jard. Bot. Buitenzorg III, 1 (1920) 383; Merrill, J. Straits Branch Roy. 
Asiat. Soc., special number (1921) 90 {'silvestris"). — Calla sylvestris Blume, 
Catalogus (1823) 62. — Scindapsus sylvestris (Blume) Kunth, Enum. pi. 3 
(1841) 64; Miq., Flora Ned. Indie 3 (1856) 187—188. - - [Scindapsus 
angustifolius Hassk., Flora 25 (2), Beibl. 1 (1842) 12, nom. Meg. — 
Rhaphidophora angustifolia (Hassk.) Schott in Bonplandia 5 (1857) 45, 
nom. Meg. — based on the type of Calla sylvestris Blume]. — Type: 
Indonesia, Java, Blume 178 (L, lecto; L, LE, isolecto, selected by Boyce, 
1999). 

Scindapsus lingulatus Hassk., Flora 25(2) Beibl. 1 (1842) 12; Schott, Prodr. 
Syst. Aroid. (1860) 378; Engl, in A. & C, DC, Monogr. Phan. 2 (1879) 248. 
— Rhaphidophora lingulata (Hassk.) Schott, Bonplandia 5 (1857) 45. — 
Monstera lingulata (Hassk.) C. Koch ex Ender, Index Aroid. (1864) 74. — 
Type: Indonesia, Java, Hasskarl s.n. (not traced, see Boyce, 1999). 

Scindapsus aruensis Engl., Bull. Soc. Tosc. Ortic. 4 (1879) 270. — Type: 
Indonesia, Melaku, Aru Islands, Gabu-lengaw, May 1873, Beccari s.n. (FI, 
holo). 



Rhaphidophora in Borneo 



67 




Figure 14. Rhaphidophora puberula Engl. 

A. flowering shoot x l / 2 ; B. leaf lamina x 7 3 ; C. venation detail x 6; D. inflorescence, spathe 
removed x 2: E. spadix detail, post-male anthesis x 6: F. spadix detail, early fruiting x 4. A & F 
from Ridley 14444: B— E from Burkill 6267. 



68 



Card. Bull. Singapore 53 (2001) 



Rhaphidophora wrayi Hook./., Fl. Brit. India 6 (1893) 544; Engl. & K. 
Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 34. — Type: Malaysia, 
Perak, Larut, Besar, April 1882, Kunstler 2939 (K, lecto, selected by Boyce, 
1999). 

Rhaphidophora gratissima Becc., Nelle Foreste di Borneo (1902) 604. — 
Rhaphidophora sylvestris var. obtusata Engl., Malesia 1 (1883) 268 — Type: 
Malaysia, Sarawak, 1 st Div., Kuching, Nov.1865, Beccari PB 952 (FI, holo; 
FI spirit 423, K, iso). 

Rhaphidophora nigrescens Ridl., J. Straits Branch Roy. Asiat. Soc. 44 (1905) 
185 ('migrescens'), synon. nov. — Type: Malaysia, Sarawak, 1 Div., Matang, 
July 1903, Ridley s.n. (SING, lecto, selected here). Ridley cites two 
conspecific syntypes, that chosen is the best specimen for identification 
purposes. The other syntype is: Malaysia, Sarawak, 19 June 1893, Bartlett 
s.n. (SING). 

Rhaphidophora motleyana Engl. & K. Krause in Engler, Pflanzenr. 37 
(IV.23B) (1908) 25; Merrill, J. Straits Branch Roy. Asiat. Soc, special 
number (1921) 89. — Type: Indonesia, Kalimantan, South Kalimantan, 
Bangarmassin, 1857—1858, Motley 741 (K, holo). 

[Scindapsus lanceolataus Miq. nom. nud. in sched. BO] 

[Pothos cucculata Zipp. nom. nud. in sched. L] 
Figure 15 

Medium to large, occasionally very large, moderately robust, leptocaul to 
semi-pachycaul homeophyllous liane to 20 m; seedling stage a non- 
skototropic shingling shoot; pre-adult plants very seldom forming small 
terrestrial colonies of appressed shingling shoots; adult shoot architecture 
comprised of greatly elongated, clinging, physiognomically unbranched, 
leafy, non-flowering stems and long, moderately elaborated, free, densely 
leafy, flowering stems later pendent under their own weight; stems smooth, 
climbing stems rectangular in cross-section, the angles often slightly winged, 
the surfaces between slightly concave, free stems rectangular to subterete 
in cross-section, green, later mid-brown, without prophyll, cataphyll and 
petiolar sheath fibre, internodes to 2.5 — 5 x 0.5 — 1 cm on clinging shoots, 
usually less stout on free shoots, separated by weakly defined, slightly 
oblique leaf scars, older stems woody; flagellate foraging stems frequent, 
often of great length. ± rectangular in cross-section; clasping roots densely 



Rhaphidophora in Borneo 



69 




Figure 15. Rhaphidophora sylvestris (Blume) Engl. 

A. portion of adult sterile stem x V,; B. stem cross-section x 1: C. flowering shoot x '/,; D. leaf 
lamina x 1: E. venation detail x 4:F. inflorescence, spathe fallen x 1: G. spadix detail, post-male 
anthesis x 4. A & B from Backer 11199: C. F & G. from de Wilde & de Wilde- Duyfies 13830: D 
& E from Afriastini 2399. 



70 



Gard. Bull. Singapore 53 (200 J) 



arising from the nodes and internodes of clinging stems, pubescent; feeding 
roots very rare, clinging, pubescent; leaves distichous on clinging and free 
shoots, those distal on flowering shoots densely so; cataphylls and prophylls 
membranous, very soon drying and falling; petiole deeply grooved adaxially, 
1 — 8.5 x 0.15 — 0.3 cm, smooth, apical and basal genicula weakly defined; 
petiolar sheath very prominent, extending to and encircling the apical 
geniculum, briefly ligulate, very soon drying and falling to leave a continuous 
conspicuous scar from the petiole base, around the top of the apical 
geniculum and back to the base; lamina entire, lanceolate-elliptic to falcate- 
lanceolate, slightly to markedly oblique, 4.5 — 32 x 1.75 — 8.5 cm, thinly 
coriaceous, upper surfaces slightly glossy, lower surfaces semi-matt, base 
subacute to briefly truncate, apex acute to slightly attenuate, with a 
prominent apiculate apicule; midrib slightly raised abaxially, slightly sunken 
adaxially; primary venation pinnate, slightly raised abaxially and adaxially; 
inter primaries parallel to primaries and only slightly less prominent, very 
slightly raised abaxially and adaxially; secondary and tertiary venation ± 
obscure in fresh material, visible as a faint reticulum in dried specimens; 
inflorescence solitary, subtended by a fully developed foliage leaf; peduncle 
compressed-cylindric, 2 — 8.5 x 0.15 — 0.5 cm; spathe cigar-shaped, stoutly 
very short-beaked. 4.5 — 7.5 x 1 — 3 cm, thinly stiff-fleshy, dull yellow, paler 
internally, soon falling at female anthesis; spadix cylindric to weakly clavate- 
cylindrical, sessile, inserted ± level on peduncle, 3 — 6 x 1 — 2.5 cm; stylar 
region mostly rhombohexagonal, 1 — 2 x 1.5—2 mm, truncate; stigma 
punctiform, raised, c. 0.25 — 0.3 mm diam.; anthers exserted at male anthesis; 
infructescence 6 — 8 x 2 — 2.5 cm. 

Distribution: Peninsular Malaysia, Sumatera, throughout Borneo, Java, Nusa 
Tenggara and Maluku. 

Habitat: Primary to disturbed secondary lowland to lower montane forest 
on a variety of substrates including sandy loam and limestone. 120 — 1800 
m altitude. 

Note: Rhaphidophora sylvestris is more variable in Borneo than in other 
parts of its range (e.g. Peninsular Malaysia, Java) but the paucity of 
collections coupled with more or less continuous variation makes it 
impossible to be confident that more than one species is involved. Of 
particular note is the occurrence of plants with coriaceous leaf laminas that 
have variously been described as Rhaphidophora gratissima Becc. (syn. R. 
sylvestris var. obtusata Engl.) and R. nigrescens Ridl.. Such plants approach 
R. conocephala in appearance (and also resemble non-Bornean R. 
crassifolia), although the flat-topped styles readily distinguish them from 



Rhaphidophora in Borneo 



71 



R. conocephala, while the juvenile shoots without conspicuously twisted 
stems separate them from R. crassifolia. Lamina thickness is not a consistent 
character, even between duplicates of the same collection. 

Other Barman specimens seen: SARAWAK. 1 Div.: G. Pueh (Mt Poi). base camp. Clemens 
& Clemens 21935 (K); Matang. Kochummen FRI 29145 (KEP, GH, K, L, SING); Kuching. 
Ridley s.n. (K): Puak. Ridley 12413 (BM, K, SING): Kuching. Tiang Bekap. G. Maja. W.L. 
Chew 714 (L. SING). SABAH. Pantai Barat: Ranau. Bt. Hampuan. Amin & Jarius SAN 
121155 (K. SAN); Mt Kinabalu. Bungal trail, Clemens & Clemens s.n. (SING): Dallas, 
Clemens & Clemens 26720 (K. SING), Clemens & Clemens 26921 (BM. BO. K. L. SING). 
Clemens & Clemens 27030 (BM. BO. K. L, SING, US), Clemens & Clemens 27312 (BM, 
BO. K. SING): Upper Kinabalu. Mementong. Mesilau Basin. Clemens & Clemens 28487 
(BM. K. L, SING); Tenompok, Clemens & Clemens 29227 (BM. BO. K. L. SING, UC); 
Ranau. Mt Kinabalu. Mesilau valley. Cockburn SAN 70108 (SAN): Kinabalu N.P., along 
Sg. Mesilau from Mesilau Camp to Mesilau Cave. Kokawa & Hotta 3995 (KYO): Kinabalu. 
Penibukan. Sg. Tahubang. Nooteboom & Aban 1582 (L). BRUNEI DARUSSALAM. Belait: 
Sg. Keduan, off Sg. Belait. Forman 1174 (BRUN. K). KALIMANTAN. Central Kalimantan: 
Sintang. HPH km 70. west camp off main (new) logging road. Church et al. 985 (BO. GH. 
K), •Borneo'. Korthals 153, Korthals s.n. (L). 

Acknowledgements 

Thanks are due to Linda Gurr for skillfully executing the illustrations that 
accompany this article. Thanks also to Dr Alistair Hay (NSW) who critically 
reviewed the manuscript and offered much useful advice. 

References 

Boyce, P.C. 1998. The genus Epipremnum Schott (Araceae — 
Monsteroideae — Monstereae) in west and central Malesia. Blumea. 43: 
183—213. 

Boyce. P.C. 1999. The genus Rhaphidophora Hassk. (Araceae- 
Monsteroideae-Monstereae) in Peninsular Malaysia, and Singapore. 
Gardens Bulletin Singapore. 51: 183 — 256. 

Boyce. P.C. 2000a. The genus Rhaphidophora Hassk. (Araceae- 
Monsteroideae-Monstereae) in the Southern and Western Indonesian 
Archipelago. Gardens Bulletin Singapore. 52: 101 — 183. 

Boyce, P.C. 2000b. The genus Rhaphidophora Hassk. (Araceae- 
Monsteroideae-Monstereae) in the Philippines. Gardens Bulletin 
Singapore. 53: 213 — 256. 



72 



Card. Bull. Singapore 53 (2001) 



Boyce, P.C. and J. Bogner. 2000. An account of neotenic species of 
Rhaphidophora Hassk. (Araceae-Monsteroideae-Monstereae) in New 
Guinea and Australia. Gardens Bulletin Singapore. 52: 89 — 100. 

Boyce, P.C. 2001. The genus Rhaphidophora Hassk. (Araceae- 
Monsteroideae- Monstereae) in New Guinea, Australia and the Tropical 
Western Pacific. Gardens Bulletin Singapore. 53: 75 — 183. 

Engler, A. and K. Krause 1908. Araceae-Monsteroideae. In A. Engler (ed.). 
Das Pflanzenreich 37 (IV.23B) 4 — 139. Engelman, Leipzig. 

Mayo, S.J., J. Bogner and P.C. Boyce. 1997. Genera of Araceae. Royal 
Botanic Gardens, Kew, U.K. 

Schimper, A.F.W. 1903. Guilds in Plant-Geography upon a Physiological 
Basis. Pt 2, ch. 2: 192 — 206. (English translation by W.R. Fisher, revised 
and edited by P. Groom & I.B. Balfour.) Clarendon Press, Oxford, 
U.K.. 

Stearn, W.T. 1992. Botanical Latin, 4 th ed., pp. xiv + 546. David & Charles, 
Newton Abbot & London, U.K. 

Strong, D.R. & Ray, T.S. 1975. Host tree location behavior of a tropical 
vine (Monstera gigantea) by skototropism. Science. 190: 804—806. 



Index Exsiccatorum 

beccarii = 1 latevaginata = 8 

conocephala = 2 lobbii = 9 

cylindrosperma = 3 megasperma = 10 

elliptica = 4 minor = 11 

elliptifolia = 5 puberula = 12 

foraminifera - 6 sylvestris = 13 
korthalsii = 7 



Aban & Petrus SAN 90675 = 1; Afriastini 116 = 9; Ahmad 15 = 12, 64 = 7; 
Ambri & Arifin 331 = 7, 354 = 3; Amdjah 7, 32. 95 7, 1084 = 11; Amin & 
Jarius SAN 121155 = 13; Amin et al. SAN 94681, SAN 97483 = 1; Ampuria 
SAN 32773 = 7; Anderson S 3084, 8007, 8032, 14547, S 23496 = 9,S 31956 = 
7; Argent et al. 93-138 = 9; 887 = 12; 9136 = 7; Ashton 154 = 7; 12626 = 12; S 
19585 = 11 



Beccari PB 833 = 1; PB 952 = 13; PB 1977, PB 2314, PB 2714 = 7; Bogner 
1346, 1393, 1507, 1504, 1562 = 1; Boyce 235 = 6; 245 - 1; 312 - 7; 345 = 3; 



Rhaphidophora in Borneo 



73 



396 = 1: 402 = 8:719 = 1: 721 = 10: 722 = 6: 728, 1389 = 8: 1391. 1400. 1414 
= 6: 1421. 1443 = 7: Brooke 8409, 9459 = 1: 8998 = 7: 10729 = 1: Brooks 
1016 = 11; Brunig S 12093 = 7: Burbidge sub \ eitch no. 215 = 7: Bum 
12938. 13001 = 1; Bum & Martin 4926 = 7: Buwalda 7917 = 11 

Chai S 31999 = 12: S. 39439 = 1: Chew 714 = 13: Church et al. 985 = 13: 
1573 = 3: 1668 = 10: Clemens 10142, 10772 = 12: Clemens & Clemens 21929 
= 4: 21934 = 1: 21935 = 13: 26405 = 7: 26453 = 12: 26495 = 7: 26720 = 13; 
26781. 26876. 26900 = 12; 26921. 27030. 27312, 28487 = 13: 28813, 29146, 
29146a = 7; 29227 = 13: 37/50. 34298 = 7; Cockburn SAN 70108 = 13: Croar 
53775 = 6: 53737 = 7: 53782 - 6: 53186 = 4: Cuadra A 2237 = 7 

Dewol & Harum SAN 89922 = 7: Dewol & Meijer SAN 131653 = 8: Dewol 
et al. SAN 118075 = 1: J.Dransfield 6302= 7: 6917 =9 

Elmer 21593 = 12: Endert 1860, 3013 = 1 

Fid His & Sumbing SAN 113389 = 12: Forman 1174 = 13: Foxworthy 40 = 1 

Gambating SAN 94588 = 9: George et al. SAN 120521 = 9: Gibot SAN 
80977 = 7; SAN 90040 = 1; Grabowski 48 = 10 

Hallier 35. 66 = 11: 999 = 1: 7764 = 3: 1187 = 12; 7555 = 1: Haviland & Hose 
3605 = 7: Hewitt 12 = 5:40 = 7; Hirano & Hotta 41. 140. 149 = 8: 1068 = 10; 
7774 = 1; 7432 = 7: 14082 = 1; Hotta 1416 = 6: 73274 = 4: 75578 = 10: 75 777 
= 8 

Jimpin SAN 122024 = 1 

Kloss SFN 19099 = 12; Kochummen FR1 29145 = 13: Kodoh ^ Aban SAN 
82024 = 9; Kokawa & Hotta 110, 471 = 1; 637. 797 = 7: 7097 = 12: 7477 = 9: 
2455 = 7; 3089 = 12; 3995 = 13; Korthals 149 = 9; 753 = 13; Kostermans 
21125 = 7: 27733 = 9: 21725 = 7; Krispinus SAN 87213, SAN 95680 = 12: 
SAN 118444 = 1 

Lantoh SAN 87848 = 4: SAN 102053 = 1: Lim et at. LSP 766 = 4: 7272 = 8 

Madani & Saigol SAN 90180 = 12; Main (sub. Polak) 1758 = 10; 7957 = 7; 
1958 = 8; 2092 = 1; Majawat & Lasan SAN 88017= 7: Mam/r S 33623 = 11: 
5 37665 = 4; Ma/tfor 770980 = 12: Meijer 1136 = 1: 2034 = 9: 2389 = 7: 
2707 = 5; 2600 = 7: 8,4/V 38784 = 9: 44074 = 1; SAN 51617 = 11: SAN 



74 



(mi, I. Bull. Singapore 53 (2001) 



122414, SAN 122429 = 7; Meijer SAN 122736 = 11; Meijer & Dewol SAN 
131749 = 9; Meijer et al SAN 136186 = 12; Mikil SAN 38691 = 7; Mogea = 
3550 = 10; Mohtar et al. S 49245 = 7; Motley 741 = 13 

Native collector D34 = 12; 966 = 9; Nicolson 942 = 12; 1252 = 7; 1258 = 9; 
7262 = 8; 1282 = 1; 1288 = 6; 1293 = 4; 1301, 1329, 1341 = 7; 1343 = 1; 7^45 
= 4; 1359 = 9; Nieuwenhuis 97, 219 - 1; Nooteboom 4327 = 3; Nooteboom 
4120 = 10; Nooteboom & Aban 1582 = 13; Nooteboom & Chai 2216 = 7 

Othman & Munting S 55097 = 10; Othman et al S 49050 = 7 

Pflie S 16001 = 7; Po/aA: 308, 645 = 9; /We 131 = 7; Posthumus 2173 = 1; 
Po«/5e« 750 = 4; 765 = 8; 562 = 9; Purseglove 4473 = 4 

Pflflp 778 = 8; 7?/d/ey 20S9 = 10; 12413 = 13; 12414 = 7 

Sands 5979 = 9; Sidyasa PBU 650 = 3; 7777 = 11; Soepadmo & Chai S 
28148 = 7; Stone et al. SAN 85269, SAN 85310 = 7 

Tangki SAN 119596 = 1; Teysmann 8105, 11338 = 11 

Van Niel 3944, 4138 = 11; 4250 = 9; 4634 = 11; Veldkamp 7910 = 9 

Winkler 238, 1250 = 10; 3448 = 11; Wiriadinata 850 = 2; Wong 242 = 1; 7727 
= 7 

Yii S 51363 = 1 



Zainudin et al. 4537 = 10 



Gardens' Bulletin Singapore 53 (2001) 75-183. 



The Genus Rhaphidophora Hassk. (Araceae- 
Monsteroideae- Monstereae) in New Guinea, Australia 
and the Tropical Western Pacific 

P.C. Boyce 

Royal Botanic Gardens, Kew. Richmond, Surrey. TW9 3AE, U.K. 

Abstract 

An alpha-taxonomic account of Rhaphidophora in New Guinea, Australia and the tropical 
western Pacific is presented as a precursor to the forthcoming Flora Malesiana Araceae 
treatment. Thirty species are recognized, of which 12 (R. cravenschoddeana. R. cryptantha, 
R. fortis, R. gorokensis. R. guamensis. R. intonsa. R. intrusa. R. jubata, R. kokodensis, R. 
mima. R. pilosa and R. waria) are novel. Twenty new synonomies are made: R. hollrungii 
and R. iboensis = R. australasica; R. pallidivenia = R. conica; R. oreophila = R. discolor. R. 
nentii = R. geniculate: R. nutans = R. microspadix: R. buergersii. R. conferta. R. drepanophylla. 
R. obliquata and R. peekelii = R. schlechteri; R. apiculata: R. engleri [syn. R. palauensis 
Engl. & K. Krause (1921) non Koidz. (1916)]. R. forbesii. R. kanehirae and R. palauensis 
Koidz. (1916) = R. spathacea: R. graeffei and R. reineckei = R. spuria: R. ledermannii = R. 
versteegii. One species (R. dahlii Engl.) is treated as doubtful. A dichotomous key to 
species is provided. All species are illustrated. 

Contents 



Introduction 75 

Infrageneric relationships 76 

Geography and endemism 78 

Rhaphidophora 78 

Key to adult flowering Rhaphidophora in New Guinea, 

Australia and the tropical western Pacific 82 

The species 88 

Acknowledgements 179 

References 179 

Index exsiccatorum 180 

Index to species 183 



Introduction 

Rhaphidophora Hassk. (including Afrorhaphidophora Engl.; at least 3 
species in tropical Africa) comprises c. 100 species of small to large, 
occasionally enormous, root-climbing lianes (sensn Schimper, 1903), rarely 



76 



Card. Bull. Singapore 53 (200 J) 



rheophytes, distributed from tropical West Africa eastwards to the western 
Pacific (Samoa), north to southern Japan (Ryukyu Islands) and south to 
Northern Australia. Rhaphidophora is one of the largest aroid genera 
represented in tropical Asia and has several nodes of diversity; the Himalaya 
(SE Nepal to NE Vietnam, roughly 17°— 23° N), West Malesia (including 
southernmost peninsular Thailand), the Philippines, and East Malesia. 
Thirty species occur in New Guinea, Australia and the tropical Western 
Pacific region. 

The last complete revision of Rhaphidophora was that of Engler and 
Krause (1908) and a summary of the taxonomic and nomenclatural history 
of Rhaphidophora was presented in Boyce (1999). 

This is the sixth in a series of papers intended to present a complete 
alpha-taxonomy of the genus Rhaphidophora. Accounts for Peninsular 
Malaysia and Singapore (Boyce, 1999), south and west Indonesia (Boyce, 
2000a), the Philippines (Boyce, 2000b), Borneo (Boyce, 2001) and for 
neotenic species in New Guinea and Australia (Boyce & Bogner, 2000) 
have been published to date. Accounts for each of India, Sri Lanka and 
the Himalaya, Thailand and Indochina, and Africa are being prepared and 
will be published separately. 

Synonymy cited is for the species, not for the review area. The reason 
for this is that synonymous names based on types from outside the review 
area are frequently applied to specimens in local herbaria. 

Infrageneric Relationships 

Boyce (2000a) proposed eight informal morpho-taxonomic groups for the 
Rhaphidophora of Indochina and Thailand. Sunda and Wallacea, each 
group named with the earliest accepted species epithet belonging to that 
group. It should be noted that these groups are a first, pragmatic attempt 
at creating morpho-taxonomic units. Molecular work underway at 
Cambridge will result in a better, infrageneric classification. 

Of the eight groups proposed by Boyce (2000a), three occur in the 
New Guinea and Pacific region: 

1. Korthalsii Group (R. korthalsii Schott). 

2. Neoguineensis Group (R. neoguineensis Engl.. R. intrusa P.C. Boyce, 
R. mima P.C. Boyce, R. schlechteri K. Krause). 

3. Spathacea (Hollrungii) Group (R. australasica F.M. Bailey, R. 
guamensis P.C. Boyce, R. spathacea Schott, R. versteegii Engl. & K. 
Krause, R. waria P.C. Boyce). The Hollrungii Group is here renamed 
the Spathacea Group, following recognition of R. spathacea as the 
earliest described attributable species. 



Rhaphidophora in New Guinea. Australia and Western Pacific 



11 



In addition, a further eight indigenous New Guinea/Pacific groups 
are proposed and described here: 

1. Brevispathacea Group (R. brevispathacea Engl. & K. Krause. R. 
stolleana Engl. & K. Krause. perhaps R. geniculata Engl.):- Long 
pendent stems (erect and clinging in R. geniculata) with much 
abbreviated inflorescence-bearing shoots arising in the leaf axils. Spathe 
marcescent. 

2. Discolor Group (R. discolor Engl. & K. Krause. R. stenophylla K. 
Krause. R. conica Engl.. R. petrieana A. Hay):- Flowering on free 
shoots. Spadix long stipitate. Rather to markedly narrow leaves. 
(Perhaps not a good group because free shoots and stipitate spadices 
recur throughout the genus but morphologically it is quite 
homogeneous.) 

3. Fortis Group (R. fortis P.C. Boyce):- Extremely stout, clinging 
flowering shoots and short-petioled leaves. Inflorescences with 
appearance of being borne in the leaf axils. 

4. Jubata Group (R. jubara P.C. Boyce):- Robust, pachycaul. flowering 
on adherent shoots. The crested reniform seeds are unique in the 
genus. 

(Similar in appearance to R. intonsa P.C. Boyce and R. 
cravenschoddeana P.C. Boyce. which are different in not having crested 
reniform seeds.) 

5. Microspadix Group (R. microspadix K. Krause):- Long, very slender 
shoots flowering at the tip. Spathe small, marcescent. 

(May be part of the Neoguineensis Group). 

6. Okapensis Group (R. gorokensis P.C. Boyce. R. kokodensis P.C. Boyce. 
R. okapensis P.C. Boyce & Bogner. R. pilosa P.C. Boyce):- Montane 
species with stiff coriaceous leaves with prominent petiolar sheathes 
that degrade into fibres, or fall completely. Inflorescences on adherent 
or free shoots. Spathe marcescent where known (perhaps early 
marcescent in R. okapensis). 

7. Pachyphylla Group (R. cryptantha P.C. Boyce & CM. Allen. R. hayi 
P.C. Boyce & Bogner. R. pachyphylla K. Krause (+ R. parvifolia 
Alderw. - Maluku):- Defined by neotenic shingling habit with petioles 
much shorter than the lamina. 

8. Spuria Group (R. intonsa. R. cravenschoddeana. R. spuria (Schott) 
Nicolson):- Fibre-covered shoot apices, large inflorescences, often 
several together as in the Spathacea Group (q.v.) . but inflorescences 
subtended by fibrous, not chartaceous, remains. 

(Possibly heterogeneous). 



78 



Card. Bull. Singapore 53 (2001) 



Geography and Endemism 

New Guinea. Australia and the western pacific region is remarkable for 
the high number of Rhaphidophora species, coupled with very high diversity 
and a high percentage of endemics (30 species of which 29 are endemic; 
only the widespread and common R. korthalsii is not endemic to the area). 
Apart from R. guamensis (endemic to Guam) and R. petrieana (endemic 
to Australia), all the rest occur on the island of New Guinea, of which 16 
species are endemic to the island. These figures, compared with those for 
the other main nodes of diversity for the genus in tropical Asia -Borneo (5 
endemics out of a total of 13 species). Peninsular Malaysia (2 out of 15), 
Sumatera (4 out of 15) and the Philippine islands (7 out of 11) - suggest 
Rhaphidophora is a genus of Gondwana origin. 

There are no data for the conservation status of any of these endemic 
species and much remains to be done by way of field studies to ascertain 
whether any or all of these species are threatened. Nevertheless, it seems 
that several species have very restricted distributions (e.g.. R. kokodensis, 
R. gorokensis, R. juhata), others are known from a single or very few 
collections (e.g., R. pilosa, R. intrusa). 

RHAPHIDOPHORA 

Rhaphidophora Hassk., Flora 25 (2) Beibl. 1 (1842) 11; Schott, Gen. Aroid. 
(1858) 77 & Prodr. Syst. Aroid. (1860) 377—388; Miquel, Ann. Mus. Bot. 
Lugd.-Bat. 3 (1867) 81—82; Engl, in A. & C. DC, Monogr. Phan. 2 (1879) 
238—248; Engl, in Beccari, Malesia, 1 (1882) 266—272. Tab. xix 6—9, xx 
1—5; Benth. & Hook, f., Gen. PI. 3(2) (1883) 993—993; Engl. & Prantl, 
Nat. Pflanzenfam. T. 2, Ab. 3 (1889) 119—120; Engl. & Prantl, Nat. 
Pflanzenfam. Nachtr. 1 (1897) 58; K. Schum. & Lauterbach, Fl. Schutzgeb. 
Sudsee (1900) 211; Engl. & K. Krause in Engl., Pflanzenr. 37 (IV.23B) 
(1908) 17—53; Engl. & Prantl, Nat. Pflanzenfam. Nachtr. 3 (1908) 29; 
Nicolson in A.C. Sm.. Fl. Vitiensis Nova, 1 (1979) 443—445, Fig. 88; Hay 
in R.J. Johns & Hay, Students' Guide Monocot. Papua New Guinea. Part 
1 (1981) 68—72 , Fig. 29; Schott, Icones aroideae et reliquiae (IDC 
Microfilm) (1983) fiche nos. 28—31, 121; Hay, Aroids of Papua New Guinea 
(1990) 83—87, Figs. 34, 35, PI. XlVb, XV '& Telopea 5 (1993) 293—300; 
Hay et al. Checklist & botanical bibliography of the aroids of Malesia, 
Australia and the tropical western Pacific. Blumea, suppl. 8 (1995) 111 — 
127; Mayo et al. Genera Araceae (1997) 118—121. PI. 14, 109 D. - 
Scindapsus subgen. Rhaphidophora (Hassk.) Miq.. Flora Ned. Indie 3 (1856) 
185. — Type: Rhaphidophora lacera Hasskarl, nom. illeg. pro. Pothos 



Rhaphidophora in .Yen- Guinea. Australia and Western Pacific 



79 



pertusus Roxb. [= Rhaphidophora perhisa (Roxb.) Schott]. 

Scindapsas Schott subgen. Pothopsis Miq.. Flora Ned. Indie 3 (1856) 187. 
— Type: Scindapsus sylvestris (Blume) Kunth [= Rhaphidophora sylvestris 
(Blume) Engl.]. 

[Raphidophora Hassk.. Cat. Hort. Bogor. (1844) 58. orth. var.]. 

Medium-sized to very large, occasionally enormous, slender to robust, 
leptocaul or pachycaul. homeophyllous or heterophyllous, rarely neotenic. 
root-climbing lianes. very seldom clustering and rheophytic and then with 
a creeping juvenile stage: cut surfaces producing clear, odourless sticky 
juice either drying ± invisibly or coagulating into yellowish, translucent 
jelly and eventually hardening to a brittle amber-like mass: seedling stage 
mostly not observed but where known either leafy at germination and 
skototropic (see Strong & Ray. 1975) by an alternating series of congested 
leafy and elongated leafless shoots or germinating to give rise to a non- 
skototropic shingling juvenile shoot: pre-adult plants often forming modest 
to extensive terrestrial colonies of varying morphological and physiological 
form (descriptive generalisations are nearly impossible), largest terrestrial 
colonies generally occurring in places of less than optimum adult growth 
potential (e.g.. depauperate tree canopy, dry. exposed sites): adult shoot 
architecture broadly divisible into three types: i. physiognomically 
unbranched clinging non-flowering stems rooting along their entire length 
giving rise to variously elaborated free lateral lateral flowering stems, or ii. 
all stems physiognomically unbranched. clinging and flowering, or iii. 
physiognomically unbranched leader and sympodial lateral stems clinging 
but only sympodial lateral stems flowering: stems with internodes of various 
lengths separated by variously prominent leaf scars, smooth or asperous or 
densely pubescent to ramentose (the last not in the review area), older 
stems sub-woody or somewhat corky or with distinctive matt to sub-lustrous 
pale brown papery epidermis, with or without variously textured prophyll. 
cataphyll and petiolar sheath fibre either at the tips or along the newer 
sections, rarely with both cataphylls and prophylls deliquescing to black 
mucilage and later drying leaving fragmentary parchment-like remains on 
petioles, developing laminas and inflorescences: flagellate foraging stems 
occurring in some species, often exceedingly long, reaching the ground 
then rooting, variously foraging and climbing again: clasping roots sparsely 
to densely arising from the nodes and internodes. strongly adherent to 
substrate: feeding roots rare to abundant, smooth pubescent or prominently 
scaly, later often becoming woody, strongly adherent to substrate or free: 



80 



Card. Bull. Singapore 53 (2001) 



cataphylls and prophylls subcoriaceous to membranous, caducous or 
degrading or deliquescing to variously persistent, variously textured, sheaths 
and fibres, these where present variously clothing upper stem before 
eventually decaying and falling; leaves distichous or weakly spiralled, evenly 
distributed or scattered or clustered distally; petiole canaliculate to weakly 
carinate, smooth or pubescent, with variously prominent apical and basal 
genicula; petiolar sheath prominent, nearly reaching to overtopping the 
geniculum, occasionally one side greatly expanded and auriculate, especially 
in juvenile plants, at first membranous to coriaceous, then soon drying 
chartaceous completely or along the margins, sometimes degrading to untidy 
variously netted or simple fibres and later variously falling leaving a scar, 
or disintegrating marginally or completely; lamina submembranous to stiffly 
chartaceous or coriaceous, lanceolate or oblong, ± oblique, base decurrent 
to unequal or cordate, apex acute to acuminate, entire to regularly pinnatifid 
or perforated, if pinnate then divisions pinnatifid to pinnatisect (Stearn, 
1992: 324), midrib often ± naked between segments, lamina occasionally 
with small to well developed perforations adjacent to the midrib and primary 
veins, these sometimes extending to lamina margin (fenestrations then 
occasionally additional to fully developed pinnae), rarely abaxially 
pubescent when expanding, rarely strongly concolorous at maturity; midrib 
usually prominently raised abaxially and prominently sunken, sometime 
flush or rarely slightly raised adaxially; primary venation ± pinnate; 
interprimaries mostly present, sub-parallel to primaries and sometimes 
indistinguishable from them but usually less prominent and often drying 
paler, usually glabrous, occasionally pubescent with domatia in the axils of 
the primary and secondary veins; secondary venation striate to reticulate, 
variously prominent, often very difficult to distinguish from primary 
venation; tertiary venation where visible reticulate to tessellate; inflorescences 
solitary to several together, first inflorescence subtended by a (usually 
fully developed) foliage leaf and/or a very swiftly disintegrating cataphyll, 
subsequent inflorescences usually each subtended by a prophyll and 
cataphyll, more rarely by a prophyll and partially to almost fully formed 
foliage leaf, inflorescences at male anthesis naked by disintegration of 
subtending cataphyll or partially to almost completely obscured by netted 
and sheet-like fibres; peduncle terete to laterally compressed; spathe ovate 
to narrowly or broadly canoe-shaped, stoutly to rather weakly beaked, 
barely gaping to opening almost flat at male anthesis, usually caducous 
before male anthesis is complete, occasionally marcescent into the early 
stages of infructescence development, stiff to rather soft- or stoutly 
coriaceous, dirty-white, greenish, cream or yellow; spadix subglobose to 
clavate-cylindrical, cylindrical or fusiform, sessile or stipitate, often obliquely 
inserted on peduncle, tapering towards the apex; flowers bisexual, naked; 



Rhaphidophora in Sew Guinea, Australia and Western Pacific 



ovary 1- to partially 2-locular. lower part + bilaterally compressed, upper 
part cylindrical and variously angled, most often rhombohexagonal. those 
upper- and lower-most on the spadix often sterile and bereft of stigma, 
those uppermost frequently either scattered or partially fused to each other 
and forming a rudimentary appendix: ovules few to many, anatropous. 
funicle long, placentas parietal to basal, sometimes ± subaxile. partial septa 
variably intrusive: stylar region well developed, usually broader than ovary, 
usually truncate apically. rarely elongate-conic: stigma sticky at female 
male anthesis. punctiform. broadly elliptic or oblong, orientation 
circumferential or longitudinal: stamens 4 — 6: filaments strap-shaped: anthers 
usually prominently exserted from between ovaries at male male anthesis. 
rarely not exserted and pollen extruded from between ovaries, thecae 
dehiscing by a longitudinal slit: infructescence with stylar regions greatly 
enlarged, transversely dehiscent, the abscission developing at the base of 
the enlarged to massive stylar region and this shedding to expose the ovary 
cavity with the many seeds embedded in variously coloured sticky pulp: 
seeds oblong, testa thin, smooth, embryo axile. straight, endosperm copious: 
pollen dicolpate. extended monosulcate to fully zonate. ellipsoid or 
hamburger-shaped, medium-sized (mean 33 urn. range 24 — 55 um) (Mayo 
et al.. 1997). exine foveolate. subreticulate. rugulate. fossulate. scabrate. 
retiscabrate. verrucate. or psilate: chromosomes 2n = 60. 120 (42. 54. 56) 
(Mayo et al.. 1997). 

Distribution: About 100 species from tropical Africa. South and South 
East Asia. Australia and the Pacific with extensions into the subtropical 
Himalaya, southern China and the southernmost islands of Japan. 

Habitat: Usually in well drained subtropical and tropical wet. humid, or 
seasonally moderately dry primary and established secondary evergreen 
forest at low to mid-montane elevations as lianescent bole-climbers, 
lithophytes. rarely rheophytes 

Etymology: Greek rhaphis. rhaphidos (needle) and pherd (I bear): refers 
to the macroscopic (to 1cm long), needle-like unicellular trichosclereids 
present in tissues. 



82 



Gard. Bull. Singapore 53 (2001) 



Key to Adult Flowering* Rhaphidophora plants in New Guinea, 
Australia and the Tropical Western Pacific 

* juvenile vegetative phases included for species where confusion is possible 
la. Shingle climber (petiole much shorter than lamina, lamina appressed 



to substrate) 2 

lb. Not as above 6 

2a. Leaf lamina cordiform, chartaceous, base cordate, posterior lobes 
overlapping; primary venation strongly reticulate 3 



2b. Leaf lamina variously shaped (not cordiform), membranous to stiffly 
coriaceous, base slightly cordate to truncate; primary venation pinnate 

4 

3a. Primary venation conspicuously silver-grey; inflorescences arising 
from behind appressed shingling leaves, carried on abbreviated shoots 
arising from the leaf axil. Papua New Guinea (not known with 
certainty) 5. R. cryptantha 

3b. Primary venation not silver grey; plant never flowering in the 
shingling phase. Indonesian Papua, Papua New Guinea (including 
New Ireland) 29. R. versteegii (juvenile phase) 

4a. Leaf lamina membranous, with c. 3 prominent veins per side, these 
running from near the base to the upper margin or tip and crossing 
over the reticulate minor venation; plant never flowering in the 

shingling phase. Throughout region 

16. R. korthalsii (juvenile phase) 

4b. Leaf lamina coriaceous to stiffly coriaceous, without prominent veins 
running from the base to the upper margin or tip; plant fertile in the 
shingling phase 5 

5a. Leaf lamina of flowering shoots with base truncate; spadix stoutly 
cigar-shaped, shortly stipitate, inserted ± level on stipe; stipe 4 — 6 x 
3 — 3.5 mm; stigma elongated, longitudinally orientated; plant with 
disarticulating side shoots functioning as vegetative propagation units. 
Indonesian Papua, Papua New Guinea [including New Britain, New 
Ireland, Bougainville and Muyua (Woodlark) Island], and Australia 
(Eastern tropical Queensland) 11. R. hayi 

5b. Leaf lamina of flowering shoots with base acute to cuneate; spadix 
narrowly cigar-shaped, long stipitate, inserted obliquely on stipe; 
stipe 6 — 10 x 2 — 2.4 mm; stigma punctiform; plant without 



Rhaphidophora in New Guinea, Australia and Western Pacific 



83 



disarticulating side shoots. Indonesian Papua, Papua New Guinea. 
21. R. pachyphylla 

6a. Leaf lamina variously pinnately divided and/or perforated; roots 
smooth or pubescent 7 

6b. Leaf lamina entire, if pinnate then with densely ramentose-scaly 
roots 8 

7a. Leaf lamina of mature plants pinnatisect to slightly to extensively 
perforated, perforations round to rhombic, extending c. + of lamina 
width on each side of the midrib; shoot tips with prominent 
chartaceous prophyll and cataphyll remains. Indonesian Papua. Papua 
New Guinea (including New Ireland) 29. R. versteegii 

7b. Leaf lamina of mature plants pinnatisect, the pinnae often perforated 
basally; active shoot tips with sparse to copious netted fibre. 
Throughout the region 16. R. korthalsii 

8a. Inflorescences arising on clinging shoots 9 

8b. Inflorescences arising on free shoots 17 

9a. Shoot tips without variously textured cataphyll, prophyll and petiolar 
sheath fibre; lamina ovate, 2.5 — 8.5 x 0.9 — 4 cm, very stiffly 
coriaceous, base cordate to rounded and slightly notched, apex long- 
acuminate with a pronounced tubule. Papua New Guinea 

20. R. okapensis 

9b. Shoot tips with variously textured cataphyll. prophyll and petiolar 
sheath fibre; leaf laminas various, but not the above combination of 
characters 10 

10a. Leaf lamina pinnatisect; feeding roots densely ramentose-scaly. 

Throughout region 16. R. korthalsii 

10b. Leaf lamina entire; feeding roots smooth or pubescent 1 1 

11a. Spadix sessile 12 

lib. Spadix stipitate 16 

12a. Inflorescences solitary; stylar region conical, stigma not prominent. 

New Guinea and Australia (eastern tropical Queensland) 

L R. australasica 

12b. Inflorescences two or more together; stylar region truncate, stigma 

prominent 13 



84 



Card. Bull. Singapore 53 (2001) 



13a. Inflorescences subtended by netted fibre and copious sheet-like 
tissue, arising two or more sequentially between foliage leaves. Papua 
New Guinea 7. R. fortis 

13b. Inflorescences each subtended by prominent chartaceous prophyll 
and one or more large cataphylls, the whole forming a synflorescence 
not interspersed with foliage leaves 14 

14a. Prophylls and cataphylls subtending inflorescences soon degrading 
and falling; spathe caducous, 16 — 24 cm long, stiff, thin; spadix slender 
cylindrical, 13 — 21 x 1 — 3 cm, long decurrent (to 2 cm) at peduncle/ 
spathe insertion. New Britain and Bougainville 14. R. jubata 

14b. Prophylls and cataphylls subtending inflorescences drying chartaceous 
and persisting into fruit maturation; spathe marcescent, 5 — 13.5 cm 
long, stiff-fleshy, very thick-walled (up to 1 cm at tip); spadix stoutly 
cylindrical, 3 — 16.5 x 1 — 1.5 cm, inserted almost level on peduncle 
15 

15a. Leaf lamina drying pale grey-green, often (but not always) profoundly 
perforated; juvenile growth phase shingling. Indonesian Papua. Papua 
New Guinea (including New Ireland) 29. R. versteegii 

15b. Leaf lamina drying strongly bicolored, adaxially dark brown, abaxially 
bright orange-brown with copious, minute tannin cells, never 
perforate; juvenile growth phase not shingling. Palau, Indonesian 
Papua, Papua New Guinea (including Woodlark (Muyua) Island), 
Federated States of Micronesia (Yap) 25. R. spathacea 

16a. Inflorescence solitary, terminating short shoots arising singly in the 
axils of several adjacent leaves, each inflorescence subtended by a 
prophyll and several degraded, netted cataphylls; spathe ovoid- 
ellipsoid, 3.5 — 6.5 cm long; spadix ovoid-ellipsoid, 2.5 — 3.5 x 1.3 — 
1.7 cm; stylar region prominently conical, hexagonal in plan view; 
stigma punctiform. New Guinea 8. R. geniculata 

16b. Inflorescences several close together in a synflorescence, each 
subtended by a prominent chartaceous prophyll and one or more 
chartaceous cataphylls, the entire synflorescence emerging from a 
mass of chartaceous cataphylls; spathe canoe-shaped, 4 — 5.5 cm long; 
spadix ylindrical, c. 3.2 x 0.9 cm; stylar region rounded-conical, almost 
circular in plan view; stigma deeply excavated. Indonesian Papua 

(Mimika Prov.) and Papua New Guinea (East Sepik Prov.) 

29. R. waria 



Rhaphidophora in Set*- Guinea. Australia and Western Pacific 



85 



17a. Shoot tips with variously textured cataphyll. prophyll and petiolar 
sheath fibre 16 

17b. Shoot tips without cataphyll. prophyll and petiolar sheath fibre 

24 

18a. Spadix stipitate 19 

18b. Spadix sessile 20 

19a. Flowering shoots extremely abbreviated, arising from axils of leaves, 
all but obscured by netted petiolar sheath and cataphyll fibre: spathe 
ovoid-cylindric. 3 — 1 cm long, thinly coriaceous: spadix slender 
cylindrical. 2.9 — 3.4 x 1.7 — 1.9 cm: stylar region depressed centrally: 
anthers exserted at male anthesis. Indonesian Papua (Mimika Digul 
Prov. Boundary 2. R. brevispathacea 

19b. Flowering shoots elongated and leaf)", not as above: spathe canoe- 
shaped. 20 — 23 cm long, stiffly very fleshy: spadix cylindrical to 
slightly cigar-shaped. 13.5 — 19 x 2.2 — 1 cm: stylar region truncate 
with a raised rim: anthers not exserted at male anthesis. Indonesian 
Papua (Yapen). Papua New Guinea (East & West Sepik. Milne Bay 
& Morobe Prov.) 6. R. discolor 

20a. Leaf lamina narrowly lanceolate to narrowly lanceolate-elliptic, not 
exceeding 25 x 6 cm. stiffly coriaceous 21 

20b. Leaf lamina ovate to oblong-elliptic or oblong lanceolate, slightly 
oblique, frequently exceeding 30 x 10 cm. up to 76 x 32. 
submembranous. subcoriaceous. only rarely coriaceous and then 
never stiffly so 22 

21a. Cataphylls. prophylls and petiolar sheath degrading to simple robust 

fibres. Papua New Guinea (Kokoda & Port Moresby) 

15. R. kokodensis 

21b. Cataphylls. prophylls and petiolar sheath degrading to weak fibres 
and strips of soft felt-like tissue. Indonesian Papua (Kepala Burung 
- Arfak Mts) 23. R. pilosa 

22a. Leaf lamina drying strongly bicolored. adaxially mid-brown, abaxially 
pale brown: inflorescences usually several together, occasionally 
solitary each subtended by a soon-degrading membranous prophyll 
and cataphyll: spathe 9.5 — 12 (-16 cm) long. Papua New Guinea 
(Western Prov.). Solomon Islands (Guadalcanal. New Georgia. San 
Cristobal and the Santa Cruz Group). Fiji. Western and American 
Samoa 26. R. spuria 



86 



Gard. Bull. Singapore 53 (21)01) 



22b. Leaf lamina not drying bicolored or if bicolored then abaxially 
greenish brown, adaxially dark brown with dense faint dark speckling; 
inflorescence always solitary, subtended by a ± fully developed foliage 
leaf and copious netted fibre and sheet-like tissue or a i fully 
developed foliage leaf and one or more degrading papery cataphylls; 
spathe 17 — 22 long 23 

23a. Leaf lamina subcoriaceous, drying concolorous, base subacute to 
slightly decurrent; inflorescence subtended by a ± fully developed 
foliage leaf and copious netted fibre and sheet-like tissue; spathe 
17 — 20 cm long; spadix 10.5 — 18.5 x 1.7 — 2.3 cm; anthers not exserted 

at male anthesis. Papua New Guinea (Central & Morobe) 

12. R. intonsa 

.23b. Leaf lamina submembranous, drying bicolored, abaxially greenish 
brown, adaxially dark brown and densely faint dark speckled, base 
oblique, one side rounded to almost truncate, the other subacute; 
inflorescence subtended by a ± fully developed foliage leaf and one 
or more degrading papery cataphylls; spathe c. 22 cm long; spadix 
17 — 19 x 1.5 — 2 cm, yellow green; anthers long-exserted at male 
anthesis. Papua New Guinea (East Sepik, Morobe & Bougainville) 
4. R. cravenschoddeana 



24a. Spadix stipitate 25 

24b. Spadix sessile 29 

25a. Stylar region conical 26 

25b. Stylar region truncate 28 

26a. Spadix base with cochleate insertion; stigma glossy, black, at tip of 
long (c. 2 mm) slender stipe; anthers not exserted at male anthesis. 
Guam 10. R. guamensis 

26b. Spadix base tapering to almost truncate; stigma not as above; anthers 
exserted at male anthesis. Papua New Guinea (including Louisiade 
Archipelago), Australia (Queensland) 27 

27a. Petiolar sheath soon drying and eventually falling more-or-less entire 
leaving a conspicuous scar; inflorescence solitary, subtended by a 
partially developed foliage leaf and a membranous cataphyll; spathe 
6 — 8 cm long; spadix stipe c. 1 x 0.6 cm. Australia (Queensland) ... 
22. R. petrieana 

27b. Petiolar sheath persistent through to leaf fall; inflorescence not 
solitary, subtended by a ± fully developed foliage leaf; spathe 1 1 — 



Rhaphidophora in Sew Guinea. Australia and Western Pacific 



87 



13 cm long: spadix stipe 2.5 — 3 x 3 — 0.4 cm. Papua New Guinea 
(including Louisiade Archipelago) 27. R. stenophvlla 

28a. Petiolar sheath extending beyond the apical geniculum by two ligules 
adaxially and by fused extensions along the abaxial midrib for up to 
half the leaf lamina length: leaf lamina oblong-elliptic, up to 16.5 cm 
long, submembranous: spathe apex truncate and extending into a 
very long, slender beak up to l h length of entire spathe: spadix 
cigar-shaped: anthers not exserted at male anthesis. Indonesian Papua 
(Kepala Burung) 13. R. intrusa 

28b. Petiolar sheath extending beyond the apical geniculum only by two 
tiny ligules: leaf lamina, lanceolate to oblong-lanceolate, up to 34 
cm long and usually exceeding 18 cm. thinly coriaceous: spathe apex 
stoutly short-beaked, beak not exceeding % length of entire spathe: 
spadix cylindrical: anthers strongly exserted at male anthesis. Papua 
New Guinea (including Bismarck Archipelago). Solomon Islands.. 
24. R. schlechteri 

29a. Stylar region conical. Primary lateral veins drying much paler than 
abaxial leaf lamina. Indonesian Papua (Kepala Burung & Mimika 
Prow). Papua New Guinea (Muyua. New Britain) 3. R. conica 

29b. Stylar region truncate. Primary lateral veins not as above 30 

30a. Leaf lamina stiffly coriaceous, narrowly lanceolate: spathe 6 — 6.5 
cm long: spadix cylindrical. 3.3 — 3.5 x c. 0.7 cm. Papua New Guinea 

(Eastern Highlands - Goroka: Southern Highlands - Mendi) 

9. R. gorokensis 

30b. Leaf lamina membranous to thinly coriaceous, variously shaped but 
not narrowly lanceolate, or if narrowly lanceolate then not stiffly 
coriaceous and inflorescences borne on the tips of long, slender, 
pendent stems: spathe 2.5—3 cm long: spadix ovoid-globose to ovoid- 
cyhndric. 1 — 1.5 x 0.5 — 0.6 cm 31 

31a. Inflorescences arising from long, pendent shoots 32 

31b. Inflorescences arising from short, spreading shoots 33 

32a. Leaf lamina narrowly lanceolate to lanceolate falcate. 4 — 19 x 1 — 3 
cm. thinly coriaceous, weakly bicolored. adaxially dull mid-brown, 
abaxially paler brown with dense tannin cells: inflorescences borne 
at tips of very long, slender flowering stems (c. 2 mm diam. in dried 
material); spathe ovoid-ellipsoid, stoutly long-beaked. 2.5 — 3 x 0.7 — 
0.9 cm: spadix ovoid-globose to ovoid-cylindric. 1 — 1.5 x 0.5 — 0.6 



88 



(ktrd. Bull. Singapore 53 (2001) 



cm. Indonesian Papua (Mimika Prov.), Papua New Guinea (Madang 

& Morobe Prov.) 

17. R. microspadix 

32b. Leaf lamina elliptic to oblong-elliptic, 20 — 29 x 8—13 cm, 
membranous, markedly bicolored, adaxially mid-brown, abaxially 
pale orange brown; inflorescences borne on short shoots arising from 
axils of leaves along long, pendent flowering stems c. 1 cm diam. in 
dried material; spathe elongate-cylindric, stoutly short-beaked, c. 
10.2 x 1.7 cm; spadix slender cylindrical, 8 x 1.1 cm. Papua New 
Guinea (East Sepik Prov.) 28. R. stolleana 

33a. Spathe cigar-shaped, 1 — 4.5 x 0.7 — 1.2 cm, caducous; spadix 
cylindrical, 1.5 — 2.5 x 0.5 — 0.8; anthers not exserted at male anthesis. 

Papua New Guinea (Bougainville & Manus), Solomon Islands 

18. R. mima 

33b. Spathe globose to ovoid-ellipsoid, 1 — 2 x 1 — 2.5 cm, marcescent; 
spadix globose to ellipsoid-cylindrical, 1 — 1.5 x 0.7 — 1 cm; anthers 

well-exserted at male anthesis. Mainland New Guinea 

19. R. neoguineensis 

The Species 

1. Rhaphidophora australasica F.M. Bailey 

Rhaphidophora australasica F.M. Bailey, Queensland Agric. J. 1 (1897) 
452 & Fl. Queensland 5 (1902) 1697; Engl. & K. Krause in Engl., Pflanzenr. 
37 (IV.23B) (1908) 28; Williams, Native PI. Queensland 1 (1979) 252, 
unnumbered plate; Jones & Gray, Climbing PI. Australia (1988) 316, 
unnumbered plate p. 322; Hay, Telopea 5 (1993) 296—297. - - Type: 
Australia, Queensland, Cairns, Cowley s.n. (BRI, holo; K, iso). 

Rhaphidophora hollrungii Engl, in K. Schum. & Hollrung, Fl. Kais.-Wilh. 
Land (1889) 19; Engl, in Bot. Jahrb. Syst. 25 (1898) 9; K. Schum. & 
Lauterbach, Fl. Schutzgeb. Siidsee (1900) 211; Engl. & K. Krause in Engl., 
Pflanzenr. 37 (IV.23B) (1908) 44, Fig. 18, synon. nov. — Type: Papua New 
Guinea ('Kaiser Wilhelmsland*), West Sepik Prov., Augusta River, Sept. 
1897, Hollrung 746 (B, lecto; selected here). Engler cited two syntypes, the 
other (Papua New Guinea ('Kaiser Wilhelmsland"), Morobe Prov., 
Sattelberg, July 1890, Lauterbach 588) is missing from Berlin and presumed 
destroyed. 



Rhaphidophora in New Guinea, Australia and Western Pacific 



89 



Rhaphidophora iboensis K. Krause. Bot. Jahrb. Syst. 49 (1912) 93. synon. 
nov. — Type: Papua New Guinea ('Kaiser Wilhelmsland"). Madang Prov., 
Ibo Range. 17 Dec. 1908. Schlechter 18985 (B. holo). 

Figure 1 

Moderately robust, pachycaul. homeophyllous liane to 15 m: seedling stage 
not observed; pre-adult plants forming sparse terrestrial colonies: adult 
shoot architecture comprised of clinging, physiognomically unbranched. 
densely leafy flowering stems: stems smooth, later with lustrous papery 
epidermis, cataphylls and prophylls soon degrading into long, untidy fibres, 
internodes 1 — 5 x 0.5 — 1 cm. separated by prominent slightly oblique corky 
leaf scars: flagellate foraging stem absent: clasping roots densely arising 
from the nodes and internodes. minutely pubescent: feeding roots not 
observed: leaves spiro-distichous: cataphylls and prophylls chartaceous but 
soon degrading into long fibres: petiole deeply grooved. 9 — 22 x 0.25 — 0.3 
cm. smooth, apical and basal geniculum prominent and drying slightly 
darker: petiolar sheath extending to apical geniculum. short-persistent, at 
first, chartaceous, soon degrading to fibres: lamina entire, lanceolate to 
elongate-lanceolate or elongate-oblanceolate. slightly falcate, 19 — 46 x 2.5 — 
9.5 cm. subcoriaceous. base rounded to subacute, apex acute to acuminate: 
midrib raised abaxially. ± sunken adaxially: primary venation pinnate, 
slightly raised abaxially. much less-so adaxially: interprimaries sub-parallel 
to primaries, and almost indistinguishable from primary lateral veins: 
secondary venation reticulate, slightly to barely visible: inflorescence solitary, 
subtended by an incompletely developed leaf and much fibrous cataphyll 
remains; peduncle slightly robust, terete. 3.5 — 9.5 x 0.2 — 0.35 cm: spathe 
slender canoe-shaped, long beaked. 5 — 9.5 x 2 — 2.5 cm. stiff-fleshy, cream, 
caducous leaving a prominent scar; spadix cylindrical, sessile, inserted 
slightly obliquely on peduncle. 3.75 — 8 x 1 — 2 cm. cream: stylar region 
long-conical, weakly hexagonal in top view, drying longitudinally ribbed. 
1.1 — 1.2 x 1 — 1.1 mm: stigma punctiform. minutely excavated, 0.1 — 0.2 x c. 
0.3 mm: anthers exserted at male anthesis: infructescence oblong-cylindrical. 
8—9 x 1.5—2 cm. 

Distribution: New Guinea (widespread but scattered; often abundant where 
it occurs). Australia (eastern tropical Queensland). 

Habitat: Rain forest. Sea level to 760 m altitude. 

Other specimens seen: INDONESIAN PAPUA. Jayapura Prov.: Cyclop Mts. above Ifaar 
Clfar'). McKee 1877 (L). PAPUA NEW GUINEA. Morobe Prov.: Lae, White et al. NGF 
1555 (L). AUSTRALIA. Queensland: Cook District. Mossman. entrance to Mossman Gorge. 



90 



Gard. Bull. Singapore 53 (2001) 




Figure 1. Rhaphidophora australasica F.M. Bailey 

A. flowering shoot x V 2 ; B. leaf lamina x V 3 ; C. venation detail x 4; D. inflorescence, spathe 
fallen x 7 2 ; E. spadix detail, post-male anthesis x 8; F. stylar region and stigma, side view x 8. 
A-C from McKee 1877; D-F from Hollaing 746. 



Rhaphidophora in New Guinea, Australia and Western Pacific 



Blake 19756 (K): c. 20 miles NNW of Daintree. Boyland 399 (K); Along road from Atherton 
to Main Coastal Highway, along Palmerston Highway. Henrietta Falls. Croat 52614 (MO); 
Wright Creek. Lake Eacham State Park, 20 km E of Atherton. Croat 52575 (K. MO); 
S.F.R. 756. East Downey L.A.. Hyland 5608 (K. L); Wright Creek, c. 10 miles E of Atherton. 
Melville 3693 (K. L. US); Lacey's Creek. Mission Beach area. Smith 10189 (K). 

Notes. 1. Almost all New Guinea specimens previously identified as R. 
australasica (incl. R. hollrungii) belong to either R. spathacea or to one of 
several new species described elsewhere in this paper. The combination of 
solitary inflorescences arising on clinging shoots, fibrous cataphyll remains 
and conical style is diagnostic for R. australasica. 

2. The two widely separated New Guinea localities suggest a species that is 
widespread but uncommon in New Guinea. 

2. Rhaphidophora brevispathacea Engl. & K. Krause 

Rhaphidophora brevispathacea Engl. & K. Krause. Bot. Jahrb. Syst. 54 
(1916) 79; K. Krause & Alderw., Nova Guinea 14 (1924) 212. — Type: 
Indonesian Papua. Mimika/Digul Prov. boundary, Lorentz River, near 
Kloofbivak, 31 Oct. 1912, Pulle 254 (B, holo; BO, K, L. iso). 

Figure 2 

Moderate, somewhat slender, leptocaul, homeophyllous (?) liane of 
unknown ultimate height; seedling and pre-adult plants not observed; adult 
shoot architecture not completely known, but observed to be comprised of 
greatly elongated, spreading to strongly pendent, physiognomically 
unbranched, leafy, non-flowering stems giving rise to extremely short, free, 
sympodial, (foliage) leafless, flowering stems: stems smooth, climbing stems 
not observed, free stems weakly rectangular to ± terete in cross-section, 
dull yellow brown, internodes to 5 x 0.7 cm. flowering shoots extremely 
abbreviated, arising from axils of leaves and. where stems pendent then 
twisting to present inflorescences upwards, stems all but obscured by netted 
petiolar sheath and cataphyll fibre: flagellate foraging stems not observed; 
clasping roots arising singly from the nodes and internodes, slightly 
pubescent, later glabrescent; feeding roots not observed; leaves weakly 
spiralled, scattered; cataphylls and prophylls membranous, very quickly 
degrading leaving semi-persistent netted remains: petiole grooved adaxially. 
7 — 9.5 x 0.2 — 0.3 cm, smooth, with a weakly defined apical and slightly 
prominent basal geniculum: petiolar sheath slightly prominent, extending 
beyond the apical geniculum by two ligules. caducous leaving a slight 
continuous scar from the petiole base, around the top of the apical 



92 



Card. Bull. Singapore 53 (2001) 



geniculum and back to the base; lamina entire, elliptic to elliptic- or 
lanceolate-elliptic, 6.2 — 20 x 2.5 — 8.75 cm, coriaceous, upper surfaces very 
slightly glossy, drying strongly bicolorous, adaxially dark brown, abaxially 
bright orange-brown with copious, minute tannin cells, base subovate, very 
briefly decurrent, apex acute to briefly acuminate, with a minute apiculate 
tubule; midrib slightly raised abaxially, sunken adaxially; primary venation 
pinnate, slightly raised abaxially, prominent (darker veins against pale 
lamina) in dried material; interprimaries parallel to, but much less distinctive 
than, primaries, very slightly raised abaxially; secondary and tertiary venation 
± invisible in dried specimens, very weakly reticulate; inflorescence solitary, 
subtended by several reduced leaves (lamina absent) with rapidly degrading 
petiolar sheaths and netted cataphylls, the whole obscuring the peduncle; 
peduncle terete, 3 — 4 x 0.25 — 0.3 cm; spathe ovoid-cylindric, stoutly very 
short-beaked, 3 — 4 x 2 — 2.3 cm, thinly coriaceous, marcescent to fruiting, 
then abcissing basally and pushed off by developing fruits, leaving a slight 
scar; spadix slender cylindrical, stipitate, inserted more or less level on 
stipe, 2.9 — 3.4 x 1.7 — 1.9 cm: stipe slender terete, c. 3 x 1.5 mm; stylar 
region rhombohexagonal, 1.7 — 2.1 x c. 2 mm, depressed centrally; stigma 
punctiform, c. 0.4 mm diam., prominent in dried material; anthers exserted 
at male anthesis; infructescence not observed. 

Distribution: Indonesian Papua (Mimika/Digul Prov. boundary). Known 
only from the type. 

Habitat: Not recorded. 100 m altitude. 

Note: Distinctive by the long pendent stems with much abbreviated, netting- 
sheathed flowering shoots and a small, marcescent spathe. Rhaphidophora 
stolleana shares the habit of long pendent stems with abbreviated of 
inflorescence-bearing shoots arising in the leaf axils, but is readily 
distinguished by a much longer spathe (c. 10.2 cm long) and spadix (c. 8 cm 
long). To date R. brevispathacea is known only from Mimika/Digul Prov. 
and R. stolleana only from East Sepik. In the small, marcescent spathe, R. 
brevispathacea resembles R. neoguineensis\ although the latter has a quite 
different growth habit and inflorescence presentation. 

3. Rhaphidophora conica Engl. 

Rhaphidophora conica Engl., Bot. Jahrb. Syst. 1 (1881) 181. — Type: 
Indonesian Papua, Kepala Burung Prov., Ramoi, Feb. 1875, Beccari PP 
409 (FI, holo). 



Rhaphidophora in New Guinea, Australia and Western Pacific 



93 




Figure 2. Rhaphidophora hrevispathacea Engl. & K. Krause 

A. adult shoot with flowering branches x '/,; B. leaf lamina x '/,; C. venation detail x 2; D. 
inflorescence, spathe fallen x 1 V 2 ; E. spadix detail, post-male anthesis x 8; F. stylar region and 
stigma, side view x 8. All from Pulle 254. 



94 



Card. Bull. Singapore 53 (2001) 



Rhaphidophora pallidivenia Alderw., Bull. Jard. Bot. Buitenzorg III, 4 
(1922) 339, synon. nov. — Type: Indonesian Papua, Mimika Province, 
Varen (Lorentz) River, 15 June 1907, Djibda s.n., ex cult. Bogor B.G. 
Y.40. April 1921, Alderwerelt s.n. (BO. holo). 

Figure 3 

Moderately robust, semi-leptocaul, homeophyllous (?) liane to 5 m; seedling 
and pre-adult plants not observed; adult shoot architecture comprised of 
elongated, clinging, physiognomically unbranched, leafy, non-flowering 
stems and elongated, free, leafy, flowering stems; stems smooth, flexuous, 
stems ± terete in cross-section, without prophyll, cataphyll fibre, internodes 
5 — 10 x 0.5 — 1 cm, separated by weak straight leaf scars, older stems woody; 
flagellate foraging stems not observed; clasping roots arising singly from 
nodes; feeding roots solitary from nodes; leaves weakly spiro-distichous, 
moderately densely arranged; cataphylls and prophylls membranous, 
caducous; petiole grooved adaxially, 7—30 x 0.6 — 1 cm, smooth, with a 
prominent apical and basal geniculum; petiolar sheath prominent, extending 
to base or top of apical geniculum. soon drying chartaceous, then degrading 
very slightly to weak fibres before falling leaving a feeble scar; lamina 
entire, elliptic to elliptic-lanceolate, 10 — 40 x 5 — 12 cm, thinly coriaceous, 
base cuneate to acute, apex acute-acuminate, with a minute tubule; midrib 
raised abaxially, slightly sunken adaxially, drying conspicuously lighter than 
surrounding lamina; primary venation pinnate, slightly raised on both 
surfaces in dried material, drying lighter; interprimaries sub-parallel to 
primaries and only slightly less conspicuous; secondary venation reticulate, 
fine but conspicuous; inflorescence solitary, subtended by a fully developed 
foliage leaf; peduncle slightly compressed-terete, 2.5 — 10 x 0.2 — 0.35 cm; 
spathe broadly cone-shaped to ellipsoid, truncate basally at insertion on 
petiole, apex briefly to rather long-beaked, 5—12 x 2 — 4 cm, thinly fleshy, 
greenish yellow, caducous, drying pale brown, falling leaving a thin scar; 
spadix cylindrical, sessile, inserted level on peduncle, 2 — 7 x 0.7 — 1.2 cm, 
obtuse, white to yellow; stylar region rounded-rhombohexagonal, 0.9 — 1.1 
x 0.9 — 1.1 mm, slightly blunt-conical; stigma punctiform, c. 0.3 mm diam., 
rather prominent in dried material; anthers slightly exserted at male anthesis; 
infructescence not observed. 

Distribution: Indonesian Papua (Kepala Burung and Mimika Prov.), Papua 
New Guinea (Muyua, New Britain). 

Habitat: Rain forests, sometimes on coralline limestone. No altitudes 
recorded. 




Figure 3. Rhaphidophora conica Engl. 

A. flowering shoot x V 6 :B. leaf lamina x V,:C. venation detail x 3;D. portion of creeping adult 
stem x '/„; E. inflorescence x 2 /,;F. spadix detail, pre-anthesis x 8. All from Furtado s.n. 



96 



Gard. Bull Singapore 53 (2001) 



Notes: 1. Brass 28608, while much smaller than the other collections, is 
morphologically identical. 

Other specimens seen: INDONESIAN PAPUA. Mimika Prov.: Varen (Lorentz) River, 
Cult. Bogor, Y40, Furtado s.n. (SING), Nicolson 926 (US). PAPUA NEW GUINEA. 
Woodlark (Muyua) Island: Kulumadu. Brass 28608 (GH, L); West New Britain Prov.: 2 
miles northeast of Kandrian, Nicolson 1552 (K, L, US). 

4. Rhaphidophora cravenschoddeana P.C. Boyce, sp. nov. 

Rhaphidophora cravenschoddeana facile cum R. intonsa confunditur (species 
etiam cum inflorescentiis grandibus et apicibus surculorum fibrosis) sed 
fibris simplicibus neque reticulatis, lamina foliae textu tenuiore, costa 
abaxialiter prominenti, venis lateralibus primariis eis interprimariis magis 
diversis, antheris longe exsertis seorsa. — TYPUS: Papua New Guinea, 
North Solomons Prov., Bougainville, near Koniguru No.l, c. 11 miles north 
of Buin, 18 July 1963, Craven & Schodde 14 (CANB, holo; GH, K, iso). 

Figure 4 

Large, robust, pachycaul (?), homeophyllous liane to unknown ultimate 
height; seedling stage not observed; pre-adult plants consisting of sparse 
terrestrial colonies; adult shoot comprised of elongated, clinging, 
physiognomically unbranched, non-flowering stems and free lateral, leafy 
flowering stems; stems with cataphylls and prophylls degrading to long 
sparse ragged fibres and weak sheets of tissue, internodes 0.5 — 3 x 0.5 — 2 
cm, separated by prominent to very large unevenly oblique corky leaf 
scars; flagellate foraging stem and clasping roots not observed; feeding roots 
stout, arising singly from the node, densely velvety-hairy; leaves spiro- 
distichous; cataphylls and prophylls quickly degrading to long sparse ragged 
fibres and weak sheets of tissue; petiole canaliculate, 6 — 46.5 x 1.3 — 1.5 cm, 
apical geniculum very large, prominent, basal geniculum very weakly 
defined; petiolar sheath very prominent, extending to apical geniculum, 
swiftly degrading to long ragged fibres and weak sheets of tissue, later 
falling leaving a jagged scar; lamina entire, ovate to oblong-elliptic or 
oblong-ovate, strongly oblique, 8 — 53 x 6 — 26 cm, submembranous, drying 
discolorous, adaxially dark brown, densely faint dark speckled, abaxially 
greenish brown, base oblique, one side rounded to almost truncate, other 
subacute, apex subacute to acuminate or briefly attenuate; midrib very 
prominently raised abaxially, slightly sunken adaxially; primary venation 
pinnate, prominently raised abaxially, very slightly raised adaxially, drying 
paler than lamina; interprimaries parallel to primaries, much less prominent, 
slightly raised abaxially, very slightly raised adaxially; secondary venation 



Rhaphidophora in New Guinea. Australia and Western Pacific 



97 




Figure 4. Rhaphidophora cravenschoddeana P.C. Boyce 

A. part of flowering shoot x '/ : ;B. leaf lamina x V^C. venation detail x 2: D. entire inflorescence 
x V 4 ; E. spadix detail, post-male anthesis x 2V 2 ; F. portion of creeping juvenile stem x 2 / 5 ; G. 
portion of pre-adult stem x V 4 ;. A-E from Craven & Schodde 14. F & G from Croat 52760. 



98 



Card. Bull. Singapore 53 (2001) 



parallel to interprimaries and only slightly less prominent, very slightly 
raised abaxially and adaxially; tertiary venation reticulate, weakly raised 
abaxially, invisible abaxially; inflorescence solitary, subtended by a + fully 
developed foliage leaf and one or more degrading papery cataphylls; 
peduncle laterally compressed, stout, 13 — 15 x 0.7 — 1 cm; spathe slender 
canoe-shaped, 22 x 4 cm, stoutly long-beaked, caducous before male 
anthesis, falling leaving a large scar; spadix stoutly cylindrical, sessile, 
inserted more or less level on peduncle, 17 — 19 x 1.5 — 2 cm, yellow green; 
stylar region hexagonal, 1.5 — 2 x 1.2 — 1.5 mm, truncate; stigma punctiform 
to slightly ellipsoid, very slightly raised, 0.4 x 0.4 — 0.5 mm diam.; anthers 
long-exserted at male anthesis; infructescence not observed. 

Distribution: Papua New Guinea (East Sepik and Morobe Prov.), 
Bougainville. 

Habitat: Primary and secondary riverine rain forest. 100-1000 m altitude. 

Notes: 1. Easily confused with Rhaphidophora intonsa (also with large 
inflorescences and fibrous shoot tips) but separated by the simple, not 
netted, fibres, thinner-textured leaf lamina with, on the abaxial surface, a 
conspicuously raised midrib and much more pronounced differentiation 
between the primary lateral and interprimary veins, and by the long-exserted 
anthers. 

2. The specific epithet is for the collectors of the type, Lyndley Craven and 
Richard Schodde now at CANB, who made many collections in Papua 
New Guinea. 

Other specimens seen: PAPUA NEW GUINEA. East Sepik Prov.: Vicinity of Wewak. 
along trail beyond Boys Town (Reform School operated by S.V.D. Missionaries). Croat 
52760 (MO), Croat 52763A (MO): Angoram, Upper Karawari River. Domstreich 77 (L): 
Morobe Prov.: Sattelberg. Clemens 8064 (B), 2066 (B); Along road to Sankewap, 1 km in 
from Sankewap River, beginning 2.7 km south of Lutheran School, Croat 52818 (MO): 
Along Busu River, 22.6 km by road SE of Lae via road past PNG University of Technology 
and Igam road past Army Base, beginning 3.8 km from asphalt road at Army Base, Croat 
52842 (MO); North Solomons Prov.: Bougainville: Maide River gorge, lower south slopes 
of Lake Lolow crater, c. 15 miles north of Buin, Craven & Schodde 286 (CANB, GH): Near 
Barilo village, c. 6 miles north of Buin. Craven & Schodde 375 (CANB. GH, K). 



Rhaphidophora in Sew Guinea. Australia and Western Pacific 



99 



5. Rhaphidophora cryptantha P.C. Boyce & CM. Allen, sp. now 

Architectura surculi R. cryptanthae earn R. hayi ob inflorescentias in apicibus 
surculorum abbreviatorum facile depulsorum ex axillis surculorum 
scindulantium exorientium feruntur arctissime revocat. R. cryptantha 
inflorescentias post folia ferenti. forma atque colore folii laminae, spatha 
minore marcescenti. spadice ellipsoidali atque stylo stigmatico longo 
nihilominus differt. Inflorescentiis in surculis post foUa appressa scindulantia 
vectis orientibus ac aperientibus R. cryptantha distincta est ab omnibus 
speciebus ceteris neotenicus (R. hayi. R. pachyphylla et R. paniflora) in 
quibus inflorescentiae in surculis a foliis porrectis portatae sunt, atque in 
genere unica est. — TYPUS: Cultivated Fairchild Tropical Garden (acc. 
no. 74407). ex Papua New Guinea. Botanic Garden. Lae. 24 Sept. 2000. 
Boyce 1447 (FTG. holo: K. M. MO. SING. iso). 

Figures 5 & 6 

Moderate-sized, slender to somewhat robust, semi-leptocaul. homeophyllous 
neotenic liane to 3 m: seedling stage unknown: pre-adult plants shingling, 
hardly or not forming terrestrial colonies: adult shoot architecture comprised 
of clinging, physiognomically unbranched. densely leafy, sterile stems and 
very abbreviated, free, flowering stems arising from the axils and remaining 
concealed behind the leaves of the main stem: stems rectangular in cross- 
section, widest side slightly convex, smooth, dark green, without prophyll 
and cataphyll fibre but with newest parts very thin, adherent, petiolar 
sheath tissue, internodes 1 — 5 x 0.2 — 0.5 cm. separated by + straight scars: 
flagellate foraging stems rather weakly developed, usually at least partially 
leafy: clasping roots arising from the internodes. prominently pubescent, 
spreading but usually not extending much beyond the leaf span: feeding 
roots not observed (absent?): leaves distichous, shingling and ascending on 
adherent shoots, densely arranged and slightly spreading on free shoots, 
on flagella shoots leaves scattered with internodes between earning a 
prominent cataphyll of short duration: cataphylls and prophylls 
membranous, caducous: petiole very shallowly grooved. 0.75 — 1.5 x 0.3 — 
0.4 cm. smooth, apical and basal genicula barely visible: petiolar sheath 
prominent, membranous, long-ligulate. margins of ligule fused, the ligule 
extending up to 3.5 cm above base of lamina and enclosing shoot apex, 
caducous but adhering to stem: lamina broadly ovate, coriaceous, base 
cordate, apex rounded with a tiny tubule: midrib barely visible abaxially 
and adaxially: primary venation reticulate, barely visible abaxially. slightly 
raised and silver-grey adaxially: interprimaries more-or-less absent: 
secondary venation reticulate, flush abaxially. very weakly raised adaxially. 
silver-grey: inflorescence sequentially produced from a shoot arising from 



100 



Card. Bull. Singapore 53 (2001) 




Figure 5. Rhaphidophora cryptantha P.C. Boyce & C. Allen 

A. abaxial view of adult stem with two flowering branches, x : /,; B. entire flowering branch 
with inflorescence, spathe cut away to show spadix x 2. All from Boyce 1447. 



Rhaphidophora in New Guinea, Australia and Western Pacific 




Figure 6. Rhaphidophora cryptantha P.C. Boyce & C. Allen 

A. adaxial view of adult stem, x 2 / 3 ; B. adaxial view of juvenile stem, x 2 / 3 ; C. flagellate stem 
All from Boyce 1447. 



102 



Card. Bull. Singapore S3 (2001) 



behind the leaf, each subtended by a very small membranous, caducous 
prophyll and one or more very reduced leaves; peduncle terete, 0.75 — 1 x 
0.2 — 0.3 cm; spathe ellipsoid, weakly beaked, 3.5 — 4 x 1.2 — 1.5 cm, spongy- 
fleshy, yellow, inflated and gaping at female anthesis and then closing and 
drying onto developing infructescence; spadix ellipsoid, stipitate, inserted 
+ level on stipe, 2.1 — 2.2 x 0.7 — 0.8 cm, creamy white; stipe 5 — 6 x 2 — 2 
mm, white; stylar region irregularly rhombohexagonal. 2 — 2.2 x 2.1 — 2.3 
mm, weakly conical; stigma very prominently raised on a c. 1 mm stipe, 
punctiform to globose, c. 0.5 mm diam.; anthers exserted at male anthesis; 
infructescence not seen. 

Distribution: Not known with certainty. The specimen used to make the 
type was introduced into cultivation from Lae Botanic Garden and it seems 
quite possible that it originated in Papua New Guinea. The plant in 
cultivation at Kew was received from Ted Green, also ex Papua New 
Guinea. 

Habitat: Unknown. In cultivation the plant is very intolerant of cool 
conditions, suggesting that it originates from low elevations. 

Notes: 1. A species of extraordinary appearance, both in flowering in a 
neotenic state (otherwise known from R. hayi, R. parvi folia (Maluku) and 
R. pachyphylla) and especially by the inflorescences arising and opening 
on shoots carried behind the appressed shingling leaves. Often the mature 
inflorescences are completely obscured by the leaves, although occasionally 
the tip of the spathe is visible. In this flowering mode R. cryptantha is 
distinct from the neotenic species listed above (in which the inflorescences 
are carried on shoots held clear of the leaves), and unique in the genus. 

The shoot architecture of R. cryptantha most closely recalls that of 
R. hayi in that the inflorescences are borne on the tips of easily dislodged 
abbreviated shoots arising from the axils of shingling shoots. However. R. 
cryptantha differs in bearing the inflorescences behind the foliage, in the 
shape and coloration of the leaf lamina, the smaller, marcescent spathe, 
ellipsoid spadix and long style. 

2. In cultivation R. cryptantha has been confused with Monstera dubia, a 
species with a similar shingling juvenile stage and variegated leaves. The 
two are readily separable by the orientation of the leaves, those of R. 
cryptantha are ascending, those of M. dubia descending. Dissection of the 
ovary of R. cryptantha reveals two intrusive parietal placentae and numerous 
ovules, assigning it to Rhaphidophora. 



Rhaphidophora in New Guinea. Australia and Western Pacific 



103 



3. Despite the as yet unconfirmed origin of this plant, we have no hesitation 
in publishing it as a new species as it is of such distinctive appearance. 

4. The specific epithet alludes to the inflorescences that open hidden or 
barely emerging from behind the shingling, substrate-clasping leaves. 

Other specimens seen: CULTIVATED. Ex Papua New Guinea. Cult. Kew acc. no. 1983- 
4495. Green s.n. (K). 

6. Rhaphidophora discolor Engl. & K. Krause 

Rhaphidophora discolor Engl. & K. Krause. Bot. Jahrb. Syst. 54 (1916) 80. 
— Type: Papua New Guinea, East Sepik Prov., April River, 7 Sept. 1912. 
Ledermann 8569 (B & B spirit, holo). 

Rhaphidophora oreophila Engl. & K. Krause. Bot. Jahrb. Syst. 54 (1916) 
80. synon. nov. — Type: Papua New Guinea. Morobe Prov.. Erap 
(Erappenberg). 31 Oct. 1912. Ledermann 9590 (B & B spirit, holo). 

Figure 7 

Medium-sized, robust, pachycaul. homeophyllous (?) liane to 25 m: seedling 
stage and pre-adidt plants not observed: adult shoot architecture not fully- 
observed but apparently comprised of elongated, clinging, physiognomically 
unbranched. non-flowering stems and free, sympodial, leafy flowering stems: 
stems smooth, cataphylls and prophylls caducous, internodes 1 — 3 x 0.5 — 1 
cm. separated by wide but not especially prominent slightly oblique corky 
leaf scars; flagellate foraging stem, clasping roots and feeding roots not 
observed: leaves distichous to very weakly spiro-distichous: cataphylls and 
prophylls caducous: petiole deeply canaliculate. 17 — 37 x 0.4 — 1 cm. apical 
geniculum prominent, basal geniculum large but weakly defined: petiolar 
sheath prominent, extending to apical geniculum. slowly degrading to 
chartaceous strips and a very few simple fibres, then falling leaving a pale 
scar; lamina entire, ovate to oblong-lanceolate or oblong-elliptic or 
oblanceolate. slightly oblique, 18 — 47 x 9 — 16 cm. coriaceous, base rounded 
to oblique and very weakly cordate on one side and slightly decurrent on 
the other, apex acute to briefly acuminate, drying adaxially dull olive- 
green and abaxially dark brown in the one known collection: midrib 
prominently raised abaxially, sunken adaxially: primary venation pinnate, 
raised abaxially and adaxially; interprimaries sub-parallel to primaries, only 
slightly less prominent, slightly raised abaxially and adaxially: secondary 
venation reticulate, slightly raised abaxially and adaxially in dry specimens: 



104 



Card. Bull. Singapore 53 (2001) 



inflorescence solitary, subtended by a fully developed foliage leaf; peduncle 
stout, terete. 9 — 10 x 1.7 — 2 cm; spathe canoe-shaped, stoutly acuminate, 
20 — 23 x 2.5 — 2.5 cm, stiffly very fleshy, marcescent, eventually falling 
leaving a very large (c. 1.5 cm wide) scar; spadi.x cylindrical to slightly 
cigar-shaped, slightly curved, long-stipitate, inserted more or less level on 
stipe, 13.5 — 19 x 2.2 — 4 cm; stipe slender-terete, 1.5 — 3 x 0.4 — 1.5 cm; stylar 
region rhombohexagonal, 1.75 — 2.5 x 1.5 — 3 mm. truncate with a raised 
rim; stigma punctiform, flat, c. 0.4 mm diam.; anthers not exserted at male 
anthesis; infructescence cylindrical, c. 17 x 2 cm (known from fragments). 

Distribution: Indonesian Papua (Yapen), Papua New Guinea (East anf 
West Sepik, Milne Bay and Morobe Provinces). 

. Habitat: Lowland swamp forest to upper hill Araucaria forest. 20-900 m 
altitude. 

Notes: 1. A remarkable and readily identifiable species with a large, very 
long-stipitate spadix. Confusion with Rhaphidophora stenophylla (also 
flowering on free shoots and with long-stipitate spadix) is possible, although 
the latter is readily identifiable by the much narrower leaf laminas, persistent 
petiolar sheath not degrading to strips and fibre, smaller (6 — 9 x 1.5 — 2.2 
cm) spadix and long, pointed style 

2. While Nicolson 1475 seems clearly to belong here on morphology, it 
occurs in very different habitat (upper hill Araucaria forest) from that of 
the type (lowland swamp). 

3. Rhaphidophora oreophila Engl. & K. Krause is a smaller manifestation 
of this species. 

Other specimens seen: INDONESIAN PAPUA. Pulau Yapen. Kamuda. near Serui. Aet & 
Idjan s.n. (L). PAPUA NEW GUINEA. West Sepik Prov.: Telefomin, Sandaun, Hak 
Valley, contour transect on south bank of Nenem river, opposite community school. Frodin 
& Morren 3209 (K): East Sepik Prov.: Angoram. Latoma village. Wogupmeri river. Leach 
VGF 34337 (L); Morobe Prov.: Araucaria forest plantation 2 miles west of Wau. Nicolson 
1475 (B, L. K. US): Milne Bav Prov.: Raba Raba. Mavu River, near Mayu Island. Streimann 
NGF 28712 (L. US). 

7. Rhaphidophora fortis P.C. Boyce, sp. nov. 

Rhaphidophora fortis in Nova Guinea singularis est characteribus 
sequentibus concatenatis: folia breviter petiolata, surculi florentes 



Rhaphidophora in .Xew Guinea. Australia and Western Pacific 



105 




Figure 7. Rhaphidophora discolor Engl. & K. Krause 

A. flowering shoot x '/,: B. leaf lamina x '/,; C. venation detail x 4: D. inflorescence, early stages 
of fruiting, spathe fallen x */ s ] E. lower spadix detail, early fruiting x 6: F. upper spadix detail, 
early fruiting x 3. All from Streimann & Kairo NGF 39190. 



106 



Gard. Bull. Singapore 53 12001) 



adhaerentes valde robusti, inflorescentiae ex axilliis foliorum enascentes ut 
videtur. — TYPUS: Papua New Guinea, West Sepik Prov., Vanimo 
SubProv., Ossima, 27 Jan. 1969, Streimann & Kairo NGF 39190 (K, holo; 
L, LAE, CANB, US, iso). 

Figure 8 

Robust, pachycaul (?), homeophyllous (?) liane to unknown ultimate height; 
seedling stage and pre-adult plants not observed; adult shoot architecture 
not fully observed but apparently comprised of elongated, clinging, 
physiognomically unbranched, flowering stems; stems with epidermis drying 
smooth and yellowish, with cataphylls and prophylls degrading to weak 
netted fibres and sheets of chartaceous tissue at the stem tips, internodes 
1 — 2.5 x 1.5 — 2 cm, separated by weakly defined leaf scars; flagellate foraging 
stem and feeding roots not observed; clasping roots arising from nodes and 
internodes, smooth with dark chartaceous epidermis; leaves distichous; 
cataphylls and prophylls very conspicuous, extending almost to apical 
geniculum of newly emerging leaves, quickly degrading to netted fibres 
and strips of chartaceous tissue; petiole stout, broadly and shallowly 
canaliculate, 14 — 16 x 0.8 — 1 cm, apical geniculum prominent, basal 
geniculum weakly defined and obscured by cataphyll remains; petiolar sheath 
very prominent, extending to apical geniculum, but very swiftly degrading 
to netted fibres, later falling leaving a smooth, scar; lamina entire, elliptic 
to oblong-elliptic or lanceolate elliptic, slightly to markedly oblique, 24 — 
31 x 8 — 12 cm, subcoriaceous, base rounded to subacute, apex acute to 
briefly acuminate, falcate; midrib raised abaxially, sunken adaxially; primary 
venation pinnate, slightly raised abaxially and adaxially; interprimaries 
parallel to primaries, less prominent, slightly raised abaxially and adaxially; 
secondary venation tessellate-reticulate, slightly raised abaxially and 
adaxially in dry specimens; inflorescences not known but apparently arising 
two or more sequentially between foliage leaves, subtended by netted 
fibre and copious sheet-like tissue; infructscences with peduncle laterally 
compressed, c. 5 x 0.6 cm, more or less obscured by cataphyll remains; 
spathe not observed; spadix massively ovoid-cylindrical, sessile, inserted 
more or less level on peduncle, 8 — 9 x c. 3 cm; stylar region rounded- 
rhombohexagonal, c. 2.3 x 2.7, truncate; stigma punctiform, slightly raised, 
c. 0.6 mm diam.; anthers not observed. 

Distribution: Papua New Guinea (West Sepik Prov.). Known only from 
the type. 

Habitat: Lowland forest. 25 m altitude. 



Rhaphidophora in Sew Guinea. Australia and Western Pacific 



107 




Figure 8. Rhaphidophora fortis P.C. Boyce 

A. flowering shoot x V 4 : B. leaf lamina x 7 4 : C venation detail x 3: D. inflorescence, spathe 
fallen x ; : E. spadix detail, post-male anthesis x 4. F. spadix detail, earlv fruiting x 4. All from 
Nicolson 1475. 



108 



Card. Bull. Singapore 53 (2001) 



Notes: 1. An extraordinary species with extremely stout, clinging flowering 
shoots and short-petioled leaves, the whole shoot resembling a plant of 
Bornean Scindapsus beccarii Engl.. Inflorescences are borne in the leaf 
axils. It is not clear from the material available whether the inflorescence 
terminates the main axis which then reiterates, displacing the inflorescence, 
or whether the inflorescence is borne on a much abbreviated shoot arising 
from the axil. 

2. The specific epithet comes from the Latin, fortis, strong or stout, in 
allusion to the plant's relatively massive form. 

8. Rhaphidophora geniculata Engl. 

Rhaphidophora geniculata Engl., Bot. Jahrb. Syst. 25 (1898) 7; K. Schum. 
& Lauterbach, Fl. Schutzgeb. Siidsee (1900) 211; Engl. & K. Krause in 
Engl., Pflanzenr. 37 (IV.23B) (1908) 25, Fig. 7; Engl. & K. Krause, Nova 
Guinea 8 (1910) 249. — Type: Papua New Guinea, Madang Prov. (?), 
Bismarck Range, 9 Sept. 1896, Lauterbach 827 (B, neo; designated here). 
In describing R. geniculata, Engler cited three syntypes [Papua New Guinea 
('Kaiser Wilhelmsland'), Morobe Prov., Sattelberg, July 1890, Lauterbach 
616a (Bf); Papua New Guinea ("Kaiser Wilhelmsland'), Madang Prov., 
Gogol river, Nov. 1890, Lauterbach 976 (Bf) & 1058a (Bf)]. The specimen 
chosen here as the neotype was determined as R. geniculata by Engler and 
matches the protologue well. 

Rhaphidophora wentii Engl. & K. Krause, Nova Guinea 8 (1910) 248 & 
Nova Guinea 8 (1912) 805, synon. nov. — Type: Indonesian Papua, 
Noordfluss, 2 June 1907, Versteeg 1191 (B, holo; BO, L, K, iso). 

Figure 9 

Rather small, slender, semi-pachycaul homeophyllous liane to 3 m; seedling 
stage and pre-adult plants not observed; adult shoot architecture comprised 
of elongated, clinging, physiognomically unbranched, leafy, sterile stems 
and very short, adherent (always?) flowering stems arising from the axils 
of the leaves; stems smooth, terete in cross-section, drying deeply 
longitudinally sulcate; with very sparse to copious netted prophyll, cataphyll 
and petiolar sheath fibre, internodes 1 — 4 x 0.5 — 1.2 cm. separated by 
large, oblique, slightly corky leaf scars; flagellate foraging stems absent; 
clasping roots arising from the nodes and internodes, pubescent; feeding 
roots not observed; leaves distichous; cataphylls and prophylls membranous, 
quickly drying and degrading into netted fibres, these later falling; petiole 
shallowly grooved, 1.4 — 27 x 0.2 — 0.7 cm, apical geniculum long but not 



Rhaphidophora in New Guinea. Australia and Western Pacific 



109 




Figure 9. Rhaphidophora geniculata Engl. 

A. flowering shoot x '/,; B. leaf lamina x %; C. venation detail x 3; D. inflorescence, spathe 
removed x 1; E. spadix detail, pre-anthesis x 6; F. stylar region and stigma, side view x 6. All 
from Johns 9870. 



110 



Card. Bull. Singapore 53 (2001) 



especially prominent, basal geniculum barely visible to rather large 
(dependent on age), moderately prominent; petiolar sheath extending to 
just below the apical geniculum degrading into semi-persistent netted fibres, 
eventually falling; lamina entire, oblanceolate or oblong-elliptic, slightly to 
markedly oblique, oblique, 1.8 — 35 x 3 — 7 cm, thinly coriaceous to weakly 
chartaceous, usually drying conspicuously discolorous, adaxially dark olive- 
brown, abaxially pale olive-green to reddish brown, base decurrent, apex 
acute, falcate-acute, acuminate or rounded-acuminate; midrib prominently 
raised and usually darker abaxially, ± flush adaxially; primary venation 
arching-pinnate, slightly raised abaxially, almost flush adaxially; 
interprimaries sub-parallel to primaries, slightly to much less prominent, 
slightly raised; secondary venation weakly reticulate, slightly raised; tertiary 
venation reticulate-striate, barely visible; inflorescence solitary, terminating 
a short shoot arising in the axils of leaves, occasionally arising from axils of 
fallen leaves, and usually inflorescences arising from several adjacent leaves, 
and thus sections of stems carrying several inflorescences in diffuse clusters, 
each inflorescence subtended by a prophyll and several degraded, netted 
cataphylls; peduncle strongly compressed-terete, 5 — 9 x 0.3 — 0.6 cm; spathe 
ovoid-ellipsoid, base decurrent and oblique, apex rounded, slender- 
acuminate, 3.5 — 6.5 x 1.5 — 2.5 cm, white, marcescent falling at fruit 
maturation leaving a large oblique scar; spadix ovoid-ellipsoid, stipitate. 
inserted obliquely on stipe, 2.5 — 3.5 x 1.3 — 1.7 cm; stipe 2 — 8 x 1 — 2 mm; 
stylar region mostly hexagonal, 1.5 — 2 x 2.1 — 2 mm, prominently conical; 
stigma punctiform on a long (c. 2 mm) stipe, c. 0.2 — 0.35 diam.; anthers not 
exserted at male anthesis. Infructescence stout ellipsoid-cylindrical, c. 4 x 2 
cm. 

Distribution: New Guinea. Widespread but, in view of the rather few 
collections, perhaps uncommon. 

Habitat: Primary and slightly disturbed lowland to upper hill forest and 
lower montane forest, 150-790 m altitude. 

Notes: 1. In flowering on clinging shoots and by the conical style 
Rhaphidophora geniculata resembles R. australasica from which it may be 
distinguished by the ovoid-ellipsoid, stipitate spadix (cylindrical and sessile 
in R. australasica) and the lack of fibrous petiolar sheath remains. It also 
differs from that species by the inflorescences borne on short shoots arising 
from the leaf axils. In this respect it is reminiscent of R. brevispathacea and 
R. stolleana, although in these the stems are pendent and not clinging. 

2. The protologue of R. wentii states the spadix to be sessile; examination 



Rhaphidophora in New Guinea, Australia and Western Pacific 



111 



of the fertile types (B. BO. L) shows this to be incorrect: all are stipitate. 
The illustration accompanying the protologue of R. geniculate! is a good 
match for the B holotype and L isotype of R. wentii. 

Other specimen seen: INDONESIAN PAPUA. Digul Prov.: Merauke, Branderhorst 324 
(L); Mimika Prov.: Freeport Concession Area. Golf Course surrounds. Johns 9970 (BO. K. 
MAN): Kali Kopi. between Kali Kopi levee and the Kopi River. Utteridge 79 (BO. K. 
MAN). PAPUA NEW GUINEA. West Sepik Prov.: Vanimo, Vanimo hinterland. Streimann 
LA E 52964 (LAE. US): Southern Highlands Prov.: Mt Bosavi. northern side. 2 — 4 km west 
of Ludesa mission station. Jacobs 9331 (L): Wasu. NE slope of Mt Bosavi. Moi et al. 196 
(BFC. L. LAE): Morobe Prov.: vicinity of Kajabit Mission. Clemens 10662 (GH): Vicinity 
of Lae. along logging road to Busu River, from 3.8 km E of Igam road and Military Base to 
c. 6 km up the road junction. Croat 52787 (MO). Croat 52788 (MO): Left off of Igam road 
past Military Reserve. Croat 52803 (MO). 

9. Rhaphidophora gorokensis P.C. Boyce, sp. nov. 

Rhaphidophora gorokensis R. kokodensi R. okapensi, R. pilodi atque R. 
wariae similis. species omnes monticolae foliis parvis rigide coriaceis sunt. 
A R. waria atque R. okapensi in surculis liberis lateralibus florenti et spadice 
sessili prompte distinguibilis est: porro a R. waria vaginis petiolaribus plus 
minusve omnio cadentibus neque in fibris fatiscentibus. atque a R. okapensi 
forma laminae foliae (anguste lanceolate vel elliptics. basi acuta, apice 
longe acuminata tubulo minuto instructa. neque ovata. basi ovata, apice 
longe acuminata tubulo manifeste) distinguitur. A R. pilode atque R. 
kokodensi (ambo in surculis liberis lateralibus florentes) R. gorokensis 
fibra vaginae petiolaris carenti distinguitur. — TYPUS: Papua New Guinea. 
Eastern Highlands Prov.. Goroka. Marafunga logging area. Upper Asaro 
Valley. 6 Sept. 1961. Womersley & Sleumer NGF 13971 (GH. holo: L. 
LAE. iso). 

Figure 10 

Small, homeophyllous (?) liane to 12 m: seedling stage and pre-aditlt plants 
not observed: adult shoot architecture not fully observed but presumably 
comprised of elongated, clinging, physiognomically unbranched. leafy, non- 
flowering stems and (observed), free, little branching, sympodial, densely 
leafy flowering stems: stems terete in cross-section, without cataphyll. 
prophyll and petiolar sheath fibre, internodes to 1 — 4 x 0.5 — 0.8 cm, 
separated by slightly oblique, corky leaf scars: flagellate foraging stems, 
clasping roots and feeding roots unknown; leaves spiro-distichous on free 
shoots: cataphylls and prophylls chartaceous, caducous: petiole deeply 
canaliculate, 6.5 — 11 x 0.15 — 0.25 cm. smooth, genicula weak: petiolar sheath 
prominent, chartaceous. extending to apical geniculum, briefly persistent. 



112 



Gard. Bull Singapore 53 (2001) 



then partially caducous, later falling leaving an irregular, corky scar; lamina 
entire, narrowly lanceolate to elliptic, oblique, falcate, 7 — 16 x 2 — 5 cm, 
stiffly coriaceous, base acute, oblique, apex long-acuminate with a minute 
tubule; midrib raised abaxially, flush to slightly sunken adaxially; primary 
venation densely pinnate, raised abaxially and adaxially; interprimaries sub- 
parallel to primaries, indistinguishable from them; secondary and tertiary 
venation conspicuously parallel-reticulate; inflorescence solitary, subtended 
by an under-developed foliage leaf; peduncle compressed-terete, 5 — 7 x 
0.2 — 0.2.5 cm; spathe narrowly canoe-shaped, stout-acuminate, 6 — 6.5 x c. 
1.5 cm, marcescent(?); spadix cylindrical, sessile, 3.3 — 3.5 x c. 0.7 cm; stylar 
region rhombohexagonal, c. 0.9 — 1 x 0.8 — 1.1 mm, truncate; stigma 
punctiform, flush with a raised rim, c. 0.5 mm diam.; anthers not exserted 
at male anthesis; infructescence not observed. 

Distribution: Papua New Guinea (Eastern Highlands Prov. - Goroka and 
Mendi). 

Habitat: Lower montane rain forest, Fagaceae and Nothofagaceae forest 
on limestone. 1600-2440 m altitude. 

Note: Similar to Rhaphidophora kokodensis, R. okapensis, R. pilosa and R. 
waria in being a montane species with small, stiffly coriaceous leaves. It is 
readily distinguishable from R. okapensis and R. waria by flowering on 
free lateral shoots and by the sessile spadix. It can be further distinguished 
from R. waria by the petiolar sheaths falling more or less entire and not 
degrading into fibres, and from R. okapensis by the shape of the leaf 
lamina (narrowly lanceolate to elliptic, base acute, apex long-acuminate 
with a minute tubule v. ovate, base cordate, apex long acuminate with a 
pronounced tubule). From R. kokodensis and R. pilosa (both flowering on 
free lateral shoots) R. gorokensis is distinguished in lacking petiolar sheath 
fibre. 

Other specimens seen: PAPUA NEW GUINEA. Southern Highlands Prov.: Mendi, Det 
Mission, 16 km SSW of Mendi, Vinas 151 (GH, K); Eastern Highlands Prov.: Goroka, 
Collins Mill, Omahaiga River valley, Mt Otto area, Robbins 870 (L). 

10. Rhaphidophora guamensis P.C. Boyce, sp. nov. 

Dum Rhaphidophoram spathaceum simulans, R. guamensis inflorescentia 
solitaria apicem surculi liberi lateralis ferenti neque e mole prophyllorum 
chartaceorum cataphyllorumque exorienti, etiam specice robuste stipitato 
statim distinguibilis est. Praeterea, stipes stigmaticus egregie longus insignis 



Rhaphidophora in Sew Guinea. Australia and Western Pacific 



113 




Figure 10. Rhaphidophora gorokensis P.C. Boyce 

A. flowering shoot x V : : B. leaf lamina x %; C. leaf lamina x 'A; D. venation detail x 4; E. 
inflorescence x 1: F. spadix detail, pre-anthesis x 8. A & B. D-F from Womerslev & Sleitmer 
NGF 13971: C from Rabbins 870. 



114 



Card. Bull Singapore 53 (2001) 



est. — TYPUS: Guam, slopes of Mt Almagosa, Naval Magazine, near 
Fena Dam, 28 April 1962, B.C. Stone 4101 (L, holo; GUAM, US, iso). 

Figure 11 

Moderately robust, medium-sized semi-leptocaul, heterophyllous(?) liane 
to unknown ultimate height; seedling stage not observed; pre-adult plants 
forming scattered terrestrial colonies; adult shoot architecture comprised of 
elongated, clinging, physiognomically unbranched, scattered-leafy, non- 
flowering stems and free, sympodial, leafy flowering stems; stems smooth, 
mid-green, cataphylls and prophylls briefly persistent then falling leaving 
bare stems, internodes 0.5 — 3 x 0.3—1 cm, separated by prominent straight 
leaf scars; flagellate foraging stem not observed; clasping roots solitary to 
somewhat densely produced arising from the nodes and internodes adjacent 
to nodes; feeding roots not observed; leaves more-or-less distichous; 
cataphylls and prophylls drying chartaceous, briefly persistent; petiole deeply 
canaliculate, 9 — 15 x 0.2 — 0.4 cm, smooth, apical geniculum large but not 
especially prominent, basal geniculum almost invisible; petiolar sheath 
extending to base of apical geniculum, broad, chartaceous, short-persistent, 
degrading to very sparse papery strips, then falling; lamina entire, oblong- 
elliptic to oblong-lanceolate or oblanceolate, slightly oblique to markedly 
oblique, 15 — 32 x 4 — 8.5 cm, submembranous to coriaceous, base rounded 
to acute, apex acute to weakly acuminate; midrib raised abaxially, + flush 
adaxially to weakly raised abaxially; primary venation pinnate, slightly raised 
abaxially, flush adaxially, drying darker than lamina; interprimaries sub- 
parallel to primaries, much less prominent, slightly raised abaxially, barely 
visible adaxially; secondary venation reticulate, raised, especially notable 
in dry material; inflorescence solitary, subtended by a fully to partially 
developed foliage leaf and one to several degraded and soon-falling 
chartaceous cataphylls; peduncle rather stout, terete, 6—8 x 0.3 — 0.5 cm; 
spathe broadly canoe-shaped, briefly stout-beaked, 10 — 12 x 2 — 7 cm, 
spongy-fleshy, very thick-walled, pale yellow, caducous, falling leaving a 
very prominent oblique scar; stipe stoutly terete, 1 — 1.5 x 0.4 — 0.5 cm; 
spadix slender to somewhat stout-cylindrical, stipitate, cochleate at insertion 
on stipe, 7 — 9 x 1 — 1.5 cm, white at male anthesis; stylar region strongly 
conical, mostly hexagonal in top view, 0.9 — 1.2 x 1 — 1.1 mm; stigma 
punctiform on the tip of a long (c. 2 mm) stipe, 0.5 — 0.2 x c. 0.3 mm, glossy 
and black in dried material; anthers not exserted at male anthesis; 
infructescence not observed. 

Distribution: Guam. 

Habitat: Mixed forest, coconut plantation, on limestone, c. 400 m altitude. 



Rhaphidophora in New Guinea, Australia and Western Pacific 




Figure 11. Rhaphidophora guamensis P.C. Boyce 

A. flowering shoot x '/ 4 ; B. leaf x V 3 ; C. pre-adult creeping shoot x V,; D 
male anthesis x 6. All from Stone 4101. 



116 



Card. Bull Singapore 53 (2001 ) 



Note: While resembling Rhaphidophora spathacea, R. guamensis is 
immediately distinguishable by the solitary inflorescence carried on the tip 
of a free lateral shoot, and which does not emerge from a mass of 
chartaceous prophyll and cataphylls, and also by the robustly stipitate spadix. 
In addition, the remarkably long stigmatic stipe is notable. 

Other specimens seen: GUAM. Mt Lamlam summit, near lighthouse, Anderson 147 (GUAM, 
US); Almagosa Springs area. Naval Magazine, Rinehart 15568 (GUAM, US). 

11. Rhaphidophora hayi P.C. Boyce & Bogner 

Rhaphidophora hayi P.C. Boyce & Bogner, Gard. Bull. Singapore 52 (2000) 
91, fig.l. — Type: Australia, Queensland, Cooroo Lands, north Johnstone 
River, near Innisfail, Nov. 1965, Webb & Tracey 7066 (BRI, holo). 

Figure 12 

Moderate-sized, slender to somewhat robust, semi-leptocaul, homeophyllous 
neotenic liane to 5 m; seedling stage a non-skototropic shingling juvenile 
shoot; pre-adult plants forming small terrestrial colonies; adult shoot 
architecture comprised of clinging, physiognomically unbranched, mostly 
densely leafy, sterile stems and abbreviated, free, flowering stems; stems 
rectangular to terete in cross-section, widest side prominently convex, 
smooth, dark green, without prophyll and cataphyll fibre but with very 
thin, adherent, petiolar sheath tissue, internodes to 8 x 1 cm, separated by 
slight + straight scars, older stems sub-woody; flagellate foraging stems 
weakly developed, usually at least partially leafy and mostly replaced by 
short, readily disarticulating free side shoots functioning as vegetative 
propagation units; clasping roots arising from the internodes, prominently 
pubescent; feeding roots c. 3 mm diam., brown, minutely pubescent, sparsely 
lenticellate; leaves distichous, shingling and ascending on adherent shoots, 
densely arranged or slightly scattered on free shoots, scattered leaves with 
internodes between carrying a prominent cataphyll of short duration; 
cataphylls and prophylls membranous, caducous; petiole deeply grooved, 
1 — 2 x 0.2 — 0.3 cm, smooth, apical and basal genicula barely visible; petiolar 
sheath prominent, caducous but adhering to stem, membranous, ligulate, 
margins of ligule fused, the ligule extending up to 3 cm above base of 
lamina and enclosing shoot apex; lamina broadly to narrowly ovate-elliptic, 
coriaceous, base truncate to cuneate or cordate (the last not on flowering 
shoots), and briefly decurrent, apex acute with a tiny tubule; midrib 
prominently raised abaxially, slightly raised adaxially; primary venation 
densely pinnate, slightly raised abaxially, somewhat impressed adaxially; 



Rhaphidophora in New Guinea. Australia and Western Pacific 



117 




Figure 12. Rhaphidophora hayi P.C. Bovce & Bogner 

A. adult shoot with flowering branch x V,; B. leaf lamina x V 2 ; C. venation detail x 3; D. pre- 
adult climbing shoot x V 4 ; E. disarticulating side shoot x '/,; F. inflorescence x 1; G. spadix 
detail at female anthesis x 10: H. spadix detail at post-male anthesis x 10. A-C from Backer 
11199: D-H from Sands el al. 2384. 



118 



Card. Bull. Singapore 53 (2001 ) 



interprimaries sub-parallel to primaries, slightly raised on both leaf surfaces; 
secondary venation reticulate, slightly raised abaxially, + flush adaxially; 
inflorescence solitary, subtended by a membranous, caducous prophyll and 
one or more similar cataphylls; peduncle slightly laterally compressed, 2 — 
3 x 0.6 — 1 cm; spathe canoe-shaped, stoutly beaked, 5.5 — 8 x 2 — 4 cm, 
stiffly fleshy, yellow, gaping wide at female anthesis and then slowly falling 
leaving a large scar at the base of the spadix; spadix stoutly cigar-shaped, 
shortly stipitate, inserted + level on stipe, 3.5 — 6 x 1 — 1.2 cm, yellow; stipe 
4 — 6 x 3 — 3.5 mm; stylar region weakly developed, mostly irregularly 
rhombohexagonal, 1.1 — 1.3 x 1 — 1.1 mm, truncate; stigma prominently 
raised, elongated, longitudinally orientated, c. 0.3 — 0.5 x 0.2 — 0.4 mm; 
anthers not exserted at male anthesis; infructescence not seen. 

Distribution: Indonesian Papua, Papua New Guinea (including New Britain, 
New Ireland, Bougainville and Muyua (Woodlark) Island), and Australia 
(eastern tropical Queensland). 

Habitat: Primary and secondary monsoon or rain forest on coralline 
limestone and basalt. 20-600 m altitude. 

Notes: 1. While resembling Rhaphidophora pachyphylla, R. hayi is 
immediately distinguishable by the flowering shoots with broader, truncate- 
based leaf laminas, the raised, larger, elongated, longitudinally orientated 
stigmatic region, the occasional foraging shoot, and by the presence of 
free, disarticulating side shoots functioning as vegetative propagation units. 
This last character occurs also in R. cryptantha (q.v.). 

2. Very similar to R. parvifolia Alderw. (Maluku: Pulau Ternate) but 
differing by the stipitate spadix. From available material it is not possible 
to tell if R. parvifolia has the disarticulating side shoots typical of R. hayi. 

3. There are many collections of R. hayi originating from Australia (mostly 
in BRI and QRS) but all except those cited here are sterile. 

Other specimens seen: INDONESIAN PAPUA. Kepala Burung Prov.: 2 km N of Manokwari, 
Nicolson 1577 (B, K, L, US). PAPUA NEW GUINEA. Woodlark (Muyua) Island: 
Kulumadau, Brass 28831 (GH, L); Central Prov.: Sogeri Plateau, 5 - 7 miles beyond Kokoda 
Trail Monument, 30 miles east of Port Moresby, Nicolson 1431 (L. US): Milne Bay Prov.: 
Esa'ala . Normanby Island, Sewa Bay, 21 Oct. 1971, Lelean & Streimann LAE 52541 (L, 
LAE, US); North Solomons Prov.: Bougainville, Arawa, McKillup's Plantation, 6 m west of 
Kieta, Nicolson 1512 (B, K, US); West New Britain Prov.: Kandrian, along road from 
airport, Nicolson 1540 (L, US); New Ireland, Namatanai, Hans Meyer Range, above Mandih 
river, near Mandih Lake. c. 6 km WNW of Taron, 30 Oct. 1975, Sands et al. 2384 (K. K 



Rhaphidophora in Sew Guinea, Australia and Western Pacific 



1 19 



(living collection acc. no. 1975-5026. K spirit no. 63938). AUSTRALIA. Queensland: Cape 
Tribulation. Rijkers 1484 (BRI). 

12. Rhaphidophora intonsa P.C. Boyce, sp. now 

Rhaphidophora intonsa apicibus surculorum reticulo fibroso denso inclusis 
distinguitur. porro inflorescentiae grandes in surculis liberis diagnostici 
sunt. Cum R. spuria earn confundere potest, sed haec basi laminae truncata 
vel leniter cordata atque apicibus surculorum multo minus fibrosis facile 
distinguitur. Cum R. australasica earn etiam confundere potest, sed ab ilia 
R. intonsa in surculis liberis florenti et inflorescentias multo maiores (spatha 
17 — 20 cm nec 5 — 9.5 cm. spadice 10.5 — 18.5 cm nec 3.75 — 8 cm) procreanti 
differt: praetera. apex regionis styiaris truncata est. neque manifeste conica. 
— TYPUS: Papua New Guinea. Central Prov.. Boridi. 30 Sept. 1935. Carr 
14313 (SING, holo; BM, L. iso). 

Figure 13 

Medium to large, robust, pachycaul (?). homeophyllous (?) liane to 12 m: 
seedling stage and pre-adult plants not observed: adult shoot architecture 
not fully observed but apparently comprised of elongated, clinging, 
physiognomically unbranched. non-flowering stems and free lateral, leafy 
flowering stems: stems with cataphylls and prophylls degrading to dense 
ragged fibres and sheets of tissue and forming dense matting at the stem 
tips, internodes 1 — 3 x 0.5 — 1.4 cm, separated by prominent slightly oblique 
corky leaf scars: flagellate foraging stem, clasping roots and feeding roots 
not observed; leaves spiro-distichous: cataphylls and prophylls quickly 
degrading to dense ragged fibres and sheets of tissue: petiole shallowly 
canaliculate. 12 — 43 x 0.4 — 0.9 cm. apical geniculum prominent, basal 
geniculum weakly defined and obscured by fibre: petiolar sheath very 
prominent, extending to apical geniculum. swiftly degrading to copious 
netted fibres, later falling leaving a smooth, corky scar: lamina entire, 
ovate to oblong-elliptic, slightly oblique. 17 — 47 x 9 — 20 cm, subcoriaceous. 
base subacute to slightly decurrent. apex acute to briefly acuminate: midrib 
prominently raised abaxially. very slightly raised adaxially: primary venation 
pinnate, slightly raised abaxially and adaxially: interprimaries parallel to 
primaries, slightly less prominent, raised abaxially. slightly impressed 
adaxially: secondary venation reticulate, slightly raised abaxially and 
adaxially in dry specimens: inflorescence solitary, mostly subtended by a + 
fully developed foliage leaf and copious netted fibre and sheet-like tissue: 
peduncle laterally compressed to terete, 9 — 13 x 0.9 — 2 cm: spathe canoe- 
shaped. 17 — 20 x 1.5 — 2.5 cm, stoutly long-beaked, stiffly fleshy, caducous 
leaving a large scar at the base of the spadix: spadix cylindrical, slightly 



120 



Gard. Bull. Singapore 53 (2001) 



curved, sessile, inserted more or less level on peduncle, 10.5 — 18.5 x 1.7 — 
2.3 cm, cream; stylar region rhombohexagonal, 1 — 2 x 1 — 1.5 mm, truncate; 
stigma punctiform, raised at male anthesis flattened in dry material, c. 0.3 
mm diam.; anthers not exserted at male anthesis; infructescence curved- 
cylindrical, c. 17 x 2 cm. 

Distribution: Papua New Guinea (Central & Morobe Prov.). 
Habitat: Montane forest. 1290-1800 m altitude. 

Notes: 1. The densely netted fibre-encased shoot tips and large inflorescences 
are diagnostic. Nicolson 1473 is smaller and less robust than the type but 
otherwise matches very well. 

2. Confusion with R. spuria is possible although the latter is readily 
distinguished by the truncate to weakly cordate lamina base and in having 
very much less fibre at the shoot tips. Confusion with R. australasica is 
possible. Rhaphidophora intonsa differs by flowering on free shoots and in 
producing much larger inflorescences (spathe 17 — 20 cm/spadix 10.5 — 18.5 
cm vs. spathe 5 — 9.5 cm/spadix 3.75 — 8 cm in R. australasica). Additionally, 
the stylar region is truncate-topped in R. intonsa and prominently conical 
in R. australasica. 

3. The specific epithet comes from the Latin intonsa, unshaven, in allusion 
to the dense, untidy prophyll and cataphyll fibre clothing the stem tips. 

Other specimens seen: PAPUA NEW GUINEA. Morobe Prov.: Ogeramnang to Malang, 
Clemens 4637 (GH); Sambangan, Clemens 7779 (B); Edie Creek road, above Wau, Nicolson 
1473 (B, K. L, P. US). 

13. Rhaphidophora intrusa P.C. Boyce. sp. nov. 

Dum Rhaphidophoram schlechteri simulans, R. intrusa apice spathae longe 
extenso (usque ad tertiam partem longitudinis spathae toto) atque ligulis 
vaginae petiolaris valde elongatis secundum costam abaxialem usque ad 
dimidiam longitudinis laminae extensis et secum adnatis statim cognoscibilis. 
Hie character etiam, in spatione minore, in R. hayi repertus est, a qua R. 
intrusa habitu non scindulanti, petiolis longioribus, lamina foliorum tenuiore 
atque longiore differt. — TYPUS: Indonesian Papua, Kepala Burung Prov., 
Kabupaten Manokwari, Kecamatan Manokwari, Arfak Plains, close to road 
from SP & to Sg. Wariori crossing, 22 April 1994, Sands 6276 (K, holo; 
BO, MAN, iso). 



Rhaphidophora in New Guinea, Australia and Western Pacific 



121 




Figure 13. Rhaphidophora intonsa P.C. Boyce 

A. flowering shoot x V 6 ; B. leaf lamina x V 4 ; C. venation detail x 2: D. portion of clinging adult 
stem x %■ E. inflorescence, spathe fallen x '/,; F. spadix detail, male anthesis x 6" F. A^E & F 
from Can 14313; D from Nicolson 1473. 



122 



Gard. Hull. Singapore 53 (2001) 



Figure 14 

Slender, leptocaul, homeophyllous liane to 5 m; seedling and pre-adult 
plants not observed; adult shoot architecture comprised of elongated, 
clinging, physiognomically unbranched, leafy, non-flowering stems and 
somewhat lengthened mostly unbranched, free, sympodial, spreading to 
pendent, rarely ascending, leafy, flowering stems; stems smooth, flexuous, 
climbing stems + terete, free stems somewhat longitudinally flattened, 
without prophyll, cataphyll and petiolar sheath fibre, internodes 1 — 6 x 0.6 
cm on clinging shoots, those on free shoots much more slender, separated 
by weak straight to slightly oblique leaf scars, older stems woody; flagellate 
foraging stems not observed; clasping roots arising sparsely from nodes of 
clinging stems, slightly pubescent; feeding roots not observed; leaves 
distichous, somewhat scattered; cataphylls and prophylls not observed; 
petiole shallowly to rather deeply grooved adaxially, 2.5—5 x 0.1 — 0.2 cm, 
smooth, with very slight apical and basal geniculum; petiolar sheath 
prominent, chartaceous, extending beyond the apical geniculum by two 
ligules adaxially and by ligules fused along the abaxial midrib for up to 
half the leaf lamina length, sheath eventually falling leaving a continuous 
scar from the petiole base, around the top of the apical geniculum and 
back to the base and remnants along the abaxial midrib; lamina entire, 
oblong-elliptic, 6 — 16.5 x 3.5 — 6 cm, submembranous, base subovate, apex 
weakly acuminate, with a minute tubule; midrib raised abaxially, slightly 
sunken adaxially in fresh material, slightly sunken abaxially and adaxially 
in dry material; primary venation pinnate, slightly raised on both surfaces 
in dried material; interprimaries sub-parallel to, but much less distinctive 
than, primaries; secondary venation very feebly reticulate, almost invisible; 
inflorescence solitary, subtended by a fully developed foliage leaf; peduncle 
slender-terete, 3.5—4 x 0.15 cm; spathe cigar-shaped, cuneate basally at 
insertion on petiole, apex truncate and extending into a very long slender 
beak up to V3 length of entire spathe, 7 — 8 x 1.3 — 1.5 cm, thick fleshy, 
marcescent(?). later pushed off by developing infructescence(?) leaving a 
large scar; spadix cigar-shaped, stipitate, inserted level on stipe, 4.5—5.5 x 
0.9 — 1.2 cm, obtuse; stipe slender terete, c. 1 x 0.2 cm; stylar region dorso- 
ventrally compressed-rhombohexagonal, 0.9 — 1.5 x 0.85 — 1.2 mm, truncate: 
stigma punctiform, c. 0.45 mm diam., prominent in dried material; anthers 
not exserted at male anthesis; infructescence stoutly cigar-shaped, c. 5 x 1.5 
cm. 

Distribution: Indonesian Papua (Kepala Burung Prov.). Known only from 
the type. 



Rhaphidophora in Sew Guinea. Australia and Western Pacific 



123 




Figure 14. Rhaphidophora intrusa P.C. Boyce 

A. flowering shoot x : /,: B. leaf lamina x 2 / 3 ; C. inflorescence x 2 / } ; D. inflorescence, spathe 
partly removed to show spadix x z / 3 . E. spadix detail, male anthesis x 6. All from Sands 6276. 



124 



Gard. Bull. Singapore 53 (2001) 



Habitat: Partially disturbed lowland rain forest. Sea level to 20 m altitude. 

Notes: 1. While resembling Rhaphidophora schlechteri, R. intrusa is 
immediately recognizable by the long extended spathe apex (up to l h the 
length of the entire spathe) and by the greatly elongated petiolar sheath 
ligules that extend fused along the abaxial midrib for up to half the leaf 
lamina length. The latter character is also found, to a much lesser extent, 
in R. hayi, from which R. intrusa differs in the non-shingling habit, longer 
petioles and thinner, longer leaf lamina. 

2. The specific epithet comes from the Latin, intrusus, to intrude, in allusion 
to the extraordinary ligules described above. 

14. Rhaphidophora jubata P.C. Boyce, sp. nov. 

Rhaphidophora jubata species valde distincta est magnitudini grandi, 
pedunculo longo, apicibus stylaribus parvis stigmatibus prominentibus atque 
seminibus cristatis reniformibus notabilis (his adhuc Rhaphidophorae non 
perscriptis). Cum R. spuria earn confundere potest, sed haec stylares maiores 
(1.6 — 2.4 x c. 2 mm diametro) atque stigmata punctiforma vel leviter 
ellipsoidea non manifeste elevata neque siccitate nigra seminaque 
ellipsoidea ecristata habet. — TYPUS: Papua New Guinea, North Solomons 
Prov., Bougainville, path from Arawa to Korpei, 10 miles southwest of 
Kieta, 1 Nov. 1964, Nicolson 1529 (L, holo; B, K, P, US, iso). 

Figure 15 

Very robust, pachycaul, homeophyllous liane to 10 m; seedling stage not 
observed; pre-adult plants forming extensive terrestrial populations; adult 
shoot architecture comprised of elongated, clinging, physiognomically 
unbranched, densely leafy, flowering; stems smooth, with cataphylls and 
prophylls degrading into weak slightly fibrous patches and then falling, 
internodes to 4 — 13 x 1 — 3.5 cm; flagellate foraging stem not observed; 
clasping roots not observed; feeding roots arising singly from nodes, robust; 
leaves spiro-distichous; cataphylls and prophylls subcoriaceous, degrading 
to weakly fibrous patches, then falling; petiole deeply canaliculate, (6 — ) 
40 — 85 x 0.35 — 0.5 cm, smooth, apical and basal genicula very large though 
not especially prominent; petiolar sheath very prominent, extending from 
between 2 / 3 petiole length up to the apical geniculum, short-persistent, 
falling leaving thin scar; lamina entire, ovate to oblong-ovate, slightly 
oblique, 21.5 — 90 x 10.5 — 35 cm, thinly coriaceous to submembranous, 
drying pale yellow-brown to brown, base oblique, broadly rounded to 



Rhaphidophora in New Guinea, Australia and Western Pacific 



125 




Figure 15. Rhaphidophora jubata P.C. Boyce 

A. flowering shoot x l / 18 ; B. leaf lamina x C. venation detail x 2; D. inflorescence, plus lower 
peduncle x : / ; ; E.spadix detail, pre-anthesis x 6: F. mature seed, side view x 20. A from Nicolson 
1496; B-C from Nicolson 1530; D-Ffrom Nicolson 1529. 



126 



Gard. Bull. Singapore 53 (2001) 



shallowly and minutely cordate, more rarely subacute, apex rounded- 
acuminate with a small apical tubule; midrib very prominently raised and 
abaxially, ± impressed adaxially; primary venation pinnate, raised abaxially 
and adaxially; interprimaries parallel to primaries, much less prominent; 
secondary venation reticulate, hardly visible; inflorescence several together, 
each subtended by a prophyll and one or more chartaceous cataphylls, 
these soon degrading; peduncle terete, 11 — 25 x 0.3 — 2 cm; spathe slender 
canoe-shaped, stoutly attenuate-beaked, 16 — 24 x 2.5 — 3 cm, stiffly thin- 
fleshy, caducous, yellow; spadix slender cylindrical, sessile but long decurrent 
(to 2 cm) on peduncle/spathe insertion, 13 — 21 x 1 — 3 cm; stylar region, 
weakly rhombohexagonal to trapezoid, 1 — 1.2 x 1 — 1.3 mm, truncate; stigma 
punctiform, prominently raised, c. 0.25 mm diam., drying glossy black; 
anthers not exserted at male anthesis; infructescence slender cylindrical, 
14.5 — 17.5 x 2.5 cm; seed reniform, laterally compressed, conspicuously 
crested along dorsal side (i.e., the side opposed to the insertion of the 
funicle), c. 1.2 x 2.5 mm at maturity, c. 8 — 12 per fruit arranged on two 
parietal placentae. 

Distribution: Papua New Guinea (New Britain and Bougainville). 

Habitat: Primary and secondary lowland to upper hill forest, along streams 
and paths. 90-980 m altitude. 

Notes: 1. A very distinct species notable for its overall large size, long 
peduncle and small stylar tops with prominently raised stigmas. Confusion 
with Rhaphidophora spuria is possible although the latter has larger stylar 
tops (1.6 — 2.4 x c. 2 mm diam.) and punctiform to slightly ellipsoid stigmas 
that are not prominently raised and do not dry glossy black, and ellipsoid 
seeds lacking a crest. 

2. The specific epithet is derived from the Latin, jubatus, crested, in allusion 
to the crested reniform seeds of this species, a form of seed not hitherto 
recorded for Rhaphidophora. This seed form is common in Epipremnum 
Schott, which differs from Rhaphidophora by having 4 (- 6) seeds at base 
of a single intrusive parietal placenta. Furthermore, the testa in 
Rhaphidophora (including R. jubata) is brittle, while that of Epipremnum 
is tough and bony. 

Other specimen seen: PAPUA NEW GUINEA. North Solomons Prov.: Bougainville, path 
up Dakao Creek near Korpei, 11 miles southwest of Kieta, Nicolson 1530 (B, K, L, P, US); 
Pavairi, Ridsdale & Lavarack NGF 30628 (LAE, US); Kapikavi. Ridsdcde & Lavarack 
NGF 31600 (LAE. US); East New Britain Prov.: Kareeba road, 2 miles west of Kerevat, 
Nicolson 1496 (US). 



Rhaphidophora in Sew Guinea, Australia and Western Pacific 



127 



15. Rhaphidophora kokodensis P.C. Boyce, sp. nov. 

Rhaphidophora kokodensis R. australasicae similis videtur sed in surculis 
liberis lateralibus florenti. stylis apice planis. et foliis minoribus angustoribus 
magis coriaceis prompte sejuncta. Porro species montana est igitur 
altitudinaliter sejuncta. Cum R. pilode (etiam in surculis liberis lateralibus 
florenti) earn confundere potest, quamquam R. kokodensis non fibras 
aspectu coacto proprio ut in R. pilode habet. — TYPUS: Papua New 
Guinea. Central Prow. Kokoda . eastern side of Lake Myola no. 1. 23 Julv 
1974, Croft et al. LAE 61974 (GH, nolo: BRI, CANB. L. LAE, iso). 

Figure 16 

Small, homeophyllous (?) liane to unknown ultimate height: seedling stage 
and pre-adidt plants not observed: adult shoot architecture comprised of 
elongated, clinging, physiognomically unbranched. leafy, non-flowering 
stems and long, moderately elaborated, free, sympodial. densely leafy 
flowering stems: stems terete in cross-section, internodes to 1 — 4 x 0.5 — 0.8 
cm, separated by slightly oblique, prominent leaf scars: flagellate foraging 
stems, clasping roots, and feeding roots unknown: leaves spiro-distichous 
on free shoots: cataphylls and prophylls chartaceous but very soon degrading 
into long tough fibres: petiole deeply canaliculate. 5 — 16 x 0.2 — 0.25 cm. 
smooth, apical geniculum quite well defined, basal geniculum weak: petiolar 
sheath prominent, chartaceous. extending to apical geniculum. very swiftly 
degrading to fibres: lamina entire, narrowly lanceolate to narrowly 
lanceolate-elliptic, occasionally slightly falcate. 13 — 25 x 1.75 — 6 cm. stiffly 
coriaceous, base acute, apex acuminate to long-acuminate with a minute 
tubule: midrib slightly raised abaxially. slightly sunken adaxially: primary- 
venation densely pinnate, slightly raised abaxially and adaxially: 
interprimaries parallel to primaries indistinguishable from them: secondary 
and tertiary venation very faintly reticulate: inflorescence solitary, subtended 
by an underdeveloped foliage leaf and much degraded cataphyll fibre: 
peduncle compressed-terete. 7 — 12 x 0.2 — 0.3 cm: spathe narrowly cigar- 
shaped, long stout-acuminate. (-4 — )8 — 9 x c. 1.3 cm. marcescent(?): spadi.x 
cylindrical, sessile, 2 — 6 x 0.8—1 cm: stylar region rhombohexagonal, c. 
0.9 — 1 x 0.8 — 1.1 mm. truncate: stigma punctiform, very slightly raised, c. 
0.4 mm diam.: anthers not exserted at male anthesis: infructescence 
cylindrical, c. 11 x 1.3 cm. 

Distribution: Papua New Guinea (Central Prov. - Kokoda and Port 
Moresby). 



128 



Card. Bull. Singapore 53 (21)111) 



Habitat: Submontane rain forest on dark brown loam. 1500-2000 m altitude. 

Note: Superficially similar to Rhaphidophora australasica but readily 
separated by its flowering on free lateral shoots, by the flat-topped styles 
and the smaller, narrower, much more coriaceous leaves. In addition, being 
a montane species, R. kokodensis is separated altitudinally. Confusion with 
R. pilosa (also flowering on free lateral shoots) is possible, although R. 
kokodensis has petiole and shoot fibres without the distinctive felted 
appearance of those of R. pilosa. 

Other specimens seen: PAPUA NEW GUINEA. Central Prov.: Port Moresby, Boridi, Carr 
13237 (BM, K, L, SING), Carr s.n. (BM); East slope of Lake Myola No. 2, Croft & Lelean 
NGF 34561 (GH, K, L). 

16. Rhaphidophora korthalsii Schott 

Rhaphidophora korthalsii Schott, Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 129; 
Engl, in A. & C. DC, Monogr. Phan. 2 (1879) 246; Engl. & K. Krause in 
Engl., Pflanzenr. 37 (IV.23B) (1908) 49—51, Fig. 21; Alderw., Bull. Jard. 
Bot. Buitenzorg III, 4 (1922) 341; Hay, Aroids of Papua New Guinea, pi. 
XV, a — c. — Type: Indonesia, Java, P.W. Korthals s.n. (L, holo; L, P. iso). 

Pothos celatocaulis N.E. Br., Gard. Chron. 13 (1880) 200. — Rhaphidophora 
celatocaulis (N.E. Br.) Alderw., Bull. Jard. Bot. Buitenzorg III, 1 (1920) 
382 & Bull. Jard. Bot. Buitenzorg III, 4 (1922) 198. — Type: Malaysia, 
Sabah, Burbidge s.n., Hort. Veitch no. 215 (K, holo; K, iso). 

Rhaphidophora maxima Engl., Bull. Soc. Tosc. Ortic. 4 (1879) 269; Beccari, 
Malesia 1 (1882) 271, Tab. xx 1—5; K. Schum. & Lauterbach, Fl. Schutzgeb. 
Siidsee (1900) 211; Engl. & K. Krause in Engl., Pflanzenr. 37 (IV.23B) 
(1908) 48^9; K. Krause & Alderw., Nova Guinea 14 (1924) 214. — Type: 
Sarawak, G. Gading, July 1866, Beccari PB 2314 (FI, lecto, selected by 
Boyce, 1999). 

Rhaphidophora tenuis Engl., Bot. Jahrb. Syst. 1 (1881) 181; Beccari, Malesia 
1 (1882) 271—272; Engl. & K. Krause in Engl., Pflanzenr. 37 (IV.23B) 
(1908) 53. - - Types: Malaysia, Sarawak, Beccari PB 1977 (FI lecto; B 
isolecto; selected by Boyce, 1999). 

Rhaphidophora korthalsii var. angustiloba Ridl. ex Engl. & K. Krause in 
Engler, Pflanzenr. 37 (IV.23B) (1908) 49. — Type: Malaysia, Sarawak, 
Matang, July 1903, Ridley s.n. (SING, lecto; selected by Boyce, 1999). 



Rhaphidophora in New Guinea, Australia and Western Pacific 



129 




Figure 16. Rhaphidophora kokodensis P.C. Boyce 

A. flowering shoot x V 3 ; B. leaf lamina x V 2 ; C. leaf lamina x 'A; D. venation detail x 3; E. 
inflorescence, spathe fallen x 7 3 : F. spadix detail, post-male anthesis x 4. A & D from Croft et 
al. LAE 61974: B-C. E-F from Carr 13237. 



130 



Card. Bull. Singapore 53 (2001) 



Rhaphidophora copelandii Engl., Bot. Jahrb. Syst. 37 (1905) 115; Engl. & 
K. Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 49. — Type: Philippines, 
Mindanao, Davao, Mt Apo, April 1904, Copeland 1193 (B holo; PNH 
isot). 

Monstera latevaginata Engl. & K. Krause in Engl., Pflanzenr. 37 (IV.23B) 
(1908) 11. — Type: Cult. Bot. Gard. Berlin, Engler & Krause s.n. (B, holo). 

Rhaphidophora grandifolia K. Krause, Bot. Jahrb. 44, Beibl. 101 (1910) 11. 
— Type: Philippines, Negros, Negros Orientale, Dumaguete (Cuernos Mts), 
March 1908, Elmer 9464 (B, holo; E, K, L, LE, MO, PNHt iso). 

Rhaphidophora trinervia Elmer, Leafl. Philipp. Bot. 8 (1919) 3073. — Type: 
Philippines, Laguna, Los Banos (Mt Maquiling), June — July 1917, Elmer 
18057 (PNH, holof ; FI, K, L, MO, P, iso). 

Rhaphidophora ridleyi Merr., J. Str. Br. Roy. As. Soc. Special Edition 
(Enum. PI. Borneo) (1921) 90. — Rhaphidophora grandis Ridl., J. Straits 
Branch Roy. Asiat. Soc. 49 (1907) 51, nom. Meg., non Schott 1858 [India = 
R. decursiva (Roxb.) Schott]. — Type: Malaysia, Sarawak, Tambusan, Sept. 
1905, Ridley 12414 (SING, holo). 

Rhaphidophora latifolia Alderw., Bull. Jard. Bot. Buitenzorg III, 4 (1922) 
341; K. Krause & Alderw., Nova Guinea 14 (1924) 213. — Type: Indonesian 
Papua, Bonggo Range, Mamberamo, Pionierbivak, 23 July 1920, Lam 711 
(BO, holo; L, iso). 

Rhaphidophora palawanensis Merr., Philipp. J. Sci. 26 (1925) 451. — Type: 
Philippines, Palawan, Malampaya Bay, Oct. 1922, Merrill BoS 11570 (PNH, 
holot; B, K, P, iso) 

Rhaphidophora trukensis Hosok., J. Jap. Bot. 13 (1937) 195. — Type: 
Federated States of Micronesia, Chuuk (Truk) Island, near Orrip, 29 July 
1939, Hosokawa 8334 (TI , holo). 

[Epipremnum multicephalum Elmer, Leafl. Philipp. Bot. 10 (1938) 3624, 
nom. inval., descr. Angl. — Based on: Philippines, Luzon, Sorsogon, Irosin 
(Mt Bulusan), May 1916. Elmer 16061 (FI, K, L, MO, P, PNHt)]. 



Rhaphidophora in New Guinea, Australia and Western Pacific 



131 



Figures 17 & 18 

Very large, occasionally enormous, slender to rather robust, pachycaul, 
heterophyllous liane to 20 m; seedling stage a non-skototropic shingling 
juvenile shoot; pre-adult plants never forming terrestrial colonies; adult 
shoot architecture comprised of greatly elongated, clinging, physiognomically 
unbranched, densely leafy flowering stems; stems smooth, bright green, 
with sparse to copious prophyll. cataphyll and petiolar sheath fibre, 
especially at the stem tips, internodes to 15 x 3.5 cm, separated by prominent 
oblique leaf scars, older stems subwoody; flagellate foraging stems absent; 
clasping roots densely arising from the nodes and internodes, prominently 
pubescent; feeding roots abundant, adherent and free, very robust, densely 
ramentose-scaly; leaves distichous: cataphylls and prophylls membranous, 
soon drying degrading to intricately reticulate fibres, these only very slowly 
falling; petiole shallowly grooved, upper part + terete, (1 — ) 9 — 65 x 0.2 — 
1.5 cm, smooth, apical and basal genicula prominent; petiolar sheath 
prominent, membranous, strongly to slightly unequal on one side, extending 
almost to or reaching the apical geniculum, of + short-duration, soon 
degrading into persistent netted fibres, these eventually falling leaving a 
prominent, slightly corky scar; shingling lamina entire, falcate-lanceolate, 
5 — 11 x 3.5 — 6 cm, base slightly cordate, pre-adult and adult lamina 
spreading, entire, pinnatipartite, pinnatisect or pinnatifid, 10 — 44 x 14 — 94 
cm, broadly oblong-elliptic to oblong lanceolate, slightly oblique, 
membranous to chartaceous or subcoriaceous, base truncate and very briefly 
decurrent, apex acute to acuminate, individual pinnae 1 — 10 cm wide, 
frequently perforated basally adjacent to the midrib, thus appearing stilted; 
midrib very prominently raised abaxially. slightly sunken adaxially; primary 
venation pinnate, raised abaxially, somewhat impressed adaxially, 2 — 4 
primary veins per pinna; interprimaries sub-parallel to primaries, slightly 
raised abaxially, slightly impressed adaxially; secondary venation strongly 
reticulate, slightly raised; tertiary venation invisible; inflorescence solitary 
to several together, first inflorescence subtended by a membranous prophyll 
and one or more cataphylls. these swiftly degrading to netted fibres, 
subsequent inflorescences subtended by one or more swiftly degrading 
cataphylls, the whole forming a mass of developing and open inflorescences 
and developing infructescences partially concealed by persistent netted 
cataphyll and prophyll remains; peduncle slightly laterally compressed to 
terete, 6 — 26 x 1 — 1.5 cm; spathe narrowly canoe-shaped, stoutly beaked, 
10 — 30 x 3 — 5 cm, stiffly fleshy, greenish to dull yellow, gaping wide at 
female anthesis and then caducous leaving a large straight scar at the base 
of the spadix; spadix cylindrical, sessile, inserted + level on peduncle, 9 — 
26 x 1.5 — 2 cm. dull green to dirty white; stylar region rather well developed, 



132 



Gard. Hull. Singapore 53 12001) 



mostly rhombohexagonal, 1.5 — 2 x c. 2 mm, slightly conical; stigma 
punctiform to slightly elliptic, if the latter then mostly longitudinally 
orientated, c. 0.3 — 0.5 x 0.2 — 0.4 mm; anthers barely exserted at male 
anthesis; infructescence 14 — 27 x 3 — 3.5 cm, dark green ripening to dull 
orange, stylar tissue abscissing to reveal orange ovary cavity pulp. 

Distribution: Widespread in south tropical Asia from Sumatera and southern 
Thailand to Borneo and the Philippines eastwards through the tropical 
western Pacific. 

Habitat: Disturbed lowland, lower and upper hill forest primary, riverine 
or secondary forest, on basalt, granite, clay and coralline limestone. 10-800 
m altitude. 

Notes: 1. Rhaphidophora korthalsii is a very widespread and variable species, 
with an extensive synonymy. However, as with Epipremnum pinnatum 
(L.) Engl. (Boyce, 1998) there are several geographical elements that, given 
more intensive study, might warrant formal taxonomic recognition. 
Unfortunately, current herbarium material is inadequate to confirm these 
plants' status and more field observations are needed. 

2. Sterile herbarium material lacking the pre-adult stage may prove difficult 
to distinguish from the Epipremnum pinnatum. Mature leaves of 'typical' 
E. pinnatum never have more than one primary lateral vein per pinna and 
the stems of R. korthalsii lack the prominent irregular whitish longitudinal 
crests and older stems the distinctive matt to sublustrous pale brown papery 
epidermis typical of E. pinnatum. The feeding roots of R. korthalsii are 
prominently scaly while those of E. pinnatum are lenticellate-corky. The 
pre-adult stage of R. korthalsii is a shingle climber with oblong-elliptic to 
ovate slightly falcate upwards pointing leaves overlapping in the manner 
of roof tiles and with c. 3 prominent veins per side, running from near the 
base of the leaf to the upper margin or tip and crossing over the minor 
venation,. 

3. Fertile material of R. korthalsii and E. pinnatum is readily separated by 
the shape of the style apex (round v. trapezoid) and the shape and 
orientation of the stigma (± punctiform and circumferential v. strongly 
linear and longitudinal) and, if fruits are mature, by seed characters. The 
fruits of R. korthalsii each contains many small ellipsoid seeds with a brittle, 
smooth testa whereas E. pinnatum has fruits with two large, strongly curved 
seeds with a bony and ornamented testa. 



Rhaphidophora in New Guinea, Australia and Western Pacific 



133 




Figure 17. Rhaphidophora korthalsii Schort 

A. pre-adult shoot x V 4 ; B. pre-adult shingling shoot x V 4 . A from Boyce 679; B from Nicolson 



134 



Gard. Bull. Singapore 53 (2001) 



4. Confusion is possible between R. korthalsii and Amydrium zippelianum 
(Schott) Nicolson although there is a suite of characters that distinguish 
them. The leaflet tips of the Amydrium species are acute to acuminate, 
those of R. korthalsii are truncate, the petiolar sheath in R. korthalsii 
extends to the apical geniculum while in Amydrium the sheath only reaches 
to the top of the basal geniculum, the remainder of the petiole being terete 
with two sharply defined low keels running its length to merge with the 
base of the leaf lamina. The feeding roots of R. korthalsii are prominently 
scaly while those of A. zippelianum are smooth. Fruiting material of R. 
korthalsii has the stylar region abcissing to reveal a pulp cavity with 
numerous, small, ellipsoid seeds whereas A. zippelianum has one or two 
large reniform to ovoid seeds in each indehiscent fruits. 

Other specimens seen: INDONESIAN PAPUA. Kepala Burung Prov.: surroundings of 
Ayawasi, Ave 4330 (BO, L); Manokwari Prov.: Pengunungan Maoke (Nassua Mts), Docters 
van Leeuwen 10750 (BO); Wariori River, river valley west of camp between Wariori and 
Mangopi rivers, c. 11 km inland, Johns 8197 (BO, K, MAN); Fanindi. 1 km west of 
Manokwari, Nicolson 1573 (B. K. L, P, US); Arfak Mountains. Mupi Dessa, trail from 
Mupi village to G. Humibou, near S. Mupi. c. 3 km from Kali Umera, between Kali Ureda 
and the confluence of Kali Ngwes and S. Mupi, Sands 6846 (K); Mimika Prov.: Freeport 
Concession Area, along road from bridge to Kuala Kenchana, along track near left turn by 
river, Barker 145 (BO, K, MAN); Kali Kopi, between Kali Kopi levee and the Kopi River, 
Johns 9796 (BO, K, MAN); Lorentz River, near Bivak Island Pulle 66 (L); Beaufort (Van 
der Sande) River Pulle 353 (K. L); Biak Island, hill northeast of Mokmer airport, Nicolson 
1567 (B, K, L, US). PAPUA NEW GUINEA. East Sepik Prov.: vicinity of Wewak, along 
trail beyond Boys Town (Reform School operated by S.V.D. Missionaries), Croat 52763 
(MO); Lordberg, Ledermann 10319 (B spirit); Malu, Ledermann 10826 (B spirit); Central 
Prov.: Port Moresby, northeast of Manumu village. Isles & Vinas NGF 33829 (US), Isles & 
Vinas NGF 34488 (L); Manus Prov.: Manus, 1 km SW of Kabuli village on south coast in 
western Manus, Sands 2779 (K); Western Prov., Kiunga. Streimann & Womersley LAE 
51847 (L, US); Morobe Prov.: vicinity of Lae, along logging road to Busu River, from 3.8 
km E of Igam road and Military Base to c. 6 km up the road junction, Croat 52792 (MO). 
Croat 52793 (MO); c. 20 km SE of Lae. along logging road to Busu River (left off Igam 
road past Military Reserve), Croat 52800 (MO); Along Busu River, 22.6 km by road SE of 
Lae (via road past PNG University of Technology and Igam road past Army Base), beginning 
3.8 km from asphalt at Army Base, Croat 52840 (MO): Lae B.G. Croat 52849^(MO): 
Bumbu logging area, 7 miles north of Lae, Nicolson 1385 (L, US); Oomsis Creek. 20 miles 
from Lae on Bulolo road, Nicolson 1483 (B, K, L, P, US); Madang Prov.: no further data. 
Ledermann 6641 (B spirit); Lower Ramu-Atitau area, SE of Aiome Patrol Post, along 
Apenam track, east side of Tiganants River (Ioka Creek), Pullen 949 (CANB, L); North 
Solomons Prov.: Bougainville, McKillip's Arawa Plantation, 6 miles west of Kieta, Nicolson 
1520 (B, K, L, US); West New Britain Prov.: Kandrian, along road to airport, Nicolson 
1542 (B, K, L, US); East New Britain Prov.: Kareeba road, 2 miles west of Kerevat, 
Nicolson 1498, (B, K. L, P, US). FEDERATED STATES OF MICRONESIA. Chuuk 
(Truk), Toi Island, Suiyota, Takamatsu 40 (K. L); Winipwoot. C.C.Y. Wong 278 (GH, US). 
SOLOMON ISLANDS. Rennell Island: Dissing 2776 (K); Kolombangara: 2 miles NNW of 
Kuzi village. Hunt RSS 2398 (K, US); San Cristobal: Ridge between Warahito and Pegato 
river, Whitmore 6204 (K, L, US). VANUATU. Erromango: Poututu. Bourdy 228 (P); 




Figure 18. Rhaphidophora korthalsii Schott 

A. flowering shoot, leaves removed x V 2 ; B. leaf lamina x V 4 ; C. venation detail x 2: D. 
inflorescence, spathe removed x 1: E. spadix detail, post floral x 8. All from Kerr 15051. 



136 



Card. Bull. Singapore 53 (2001) 



Ipota, Bourdy 244 (K, P); Portnarvin, Rautop, Cabalion 2370 (P); vicinity of Nouankao 
Camp, Green RSNH 1267 (K); Forestry route, km 16.5, Raynal RSNH 16222 (K); Espiritu 
Santo: Bank of Achone River, opposite side from Casevaia village. Curry 939 (K); Kuvutant, 
Ludvigson 20 (L); Anatom (Aneityum): Anelgauhat Bay, Kajewski 830 (K, P), Morrison 
s.n. (K). 

17. Rhaphidophora microspadix K. Krause 

Rhaphidophora microspadix K. Krause, Bot. Jahrb. Syst. 49 (1912) 92. — 
Type: Papua New Guinea ('Kaiser WilhelmslancT), Madang Prov., Bismarck 
Range, 11 Nov. 1908, Schlechter 18678 (B, holo). 

Rhaphidophora nutans Ridl., Trans. Linn. Soc. Lond. 9 (1916) 239, synon. 
nov. — Type: Indonesian Papua, Mimika Prov., Puncak Jaya (Mt. 
Carstensz), Camp 6a, 16 Jan. 1913, Boden Kloss s.n. (BM, holo; K iso). 

Figure 19 

Small, very slender, leptocaul, homeophyllous (?) liane to unknown ultimate 
height; seedling and pre-adidt plants not observed; adult shoots comprised 
of greatly elongated, clinging, physiognomically unbranched, leafy, non- 
flowering stems and long, free, sympodial, leafy flowering stems; stems 
smooth, somewhat flexuous, stems terete in cross-section, branching little, 
growing to considerable lengths and pendent with flowering tips upturned, 
without prophyll, cataphyll and petiolar sheath fibre, internodes to 4 x c. 
0.2 cm, separated by weak slightly oblique leaf scars, older stems woody; 
flagellate foraging stems absent; clasping roots arising singly from each 
node; feeding roots not observed; leaves weakly distichous and sparsely 
arranged; cataphylls and prophylls membranous, caducous; petiole deeply 
grooved adaxially, 2 — 5 x 0.15 — 0.25 cm, smooth, with a slight apical and 
basal geniculum; petiolar sheath slightly prominent, extending to the apical 
geniculum, caducous in strips leaving a slender scar; lamina entire, narrowly 
lanceolate to lanceolate falcate, 4 — 19 x 1 — 3 cm, thinly coriaceous, drying 
adaxially dull mid-brown, abaxially paler brown with dense, well-defined 
to somewhat obscure tannin cells, base cuneate, apex long-acuminate with 
a slender prominent tubule; midrib slightly raised abaxially and adaxially; 
primary venation pinnate, very slightly raised on both surfaces; interprimaries 
sub-parallel to, but much less distinctive than, primaries, sometimes 
degrading into weakly reticulate venation, very slightly raised abaxially; 
secondary venation ± invisible in dried specimens, parallel-reticulate; 
inflorescence solitary, subtended by a fully developed foliage leaf and a 
caducous cataphyll; peduncle compressed-terete, 3 — 7.3 x 0.2 — 0.25 cm; 
spathe ovoid-ellipsoid, stoutly long-beaked and tipped with a fine tubule. 




Figure 19. Rhaphidophora microspadix K. Krause 

A. flowering shoot x V 2 ; B. leaf lamina x 1V 2 ; C. venation detail x 6; D. inflorescence, spathe 
fallen x 4. All from Boden Kloss s.n. 



138 



Card. Bull. Singapore S3 (2001 ) 



2.5 — 3 x 0.7 — 0.9 cm, marcescent until early fruiting, then falling leaving a 
large, oblique scar; spadix ovoid-globose to ovoid-cylindrical, sessile, 
inserted slightly obliquely on peduncle, 1 — 1.5 x 0.5 — 0.6 cm; stylar region 
rhombohexagonal, 1.4 — 2 x c. 2 mm, truncate; stigma punctiform, c. 0.4 
mm diam.. prominent in dried material; anthers well-exserted at male 
anthesis; infructescence ovoid-globose, c. 1.2 — 2.5 x 1 — 1.5 cm. 

Distribution: Indonesian Papua (Mimika Prov.), Papua New Guinea 
(Madang and Morobe Provinces). 

Habitat: Lower montane forest. 945-1700 m altitude. 

Notes: 1. The type specimen of Rhaphidophora microspadix has the abaxial 
leaf lamina with dense conspicuous tannin cells visible to the naked eye. 
These cells are also present on the type of R. nutans, but much less clearly 
visible (xlO lens is required). 

2. Rhaphidophora microspadix appears to be allied to R. neoguineense, 
differing in the much narrower leaf lamina with a long acuminate tip and 
in flowering habitually on long, pendent shoots. 

Other specimens seen: PAPUA NEW GUINEA. Morobe Prov.: Matap, Clemens 11151, 
41197 (GH); Gumi Divide, 25 km west of Bulolo, Kairo 757 (L. LAE). 

18. Rhaphidophora mima P.C. Boyce, sp. nov. 

Rhaphidophora mima multum cum R. neoguineensi (non in Insulas 
Solomonenses inventa) persimili confusa est, quamquam ab ea spatha 
fusiformi caduca atque spadice cylindrica (spatha globosa marcescensque 
et spadix globoso-ellipsoideus in R. neoguineensi) prompte distinguitur. — 
TYPUS: Solomon Islands, Guadalcanal, Wanderer Bay area, 23 Oct. 1968, 
Mauriasi et al. BSIP 12268 (SING, holo; BSIP, K, iso). 

Figure 20 

Slender, leptocaul, homeophyllous liane to 4 m; seedling and pre-adult 
plants not observed; adult shoot architecture not fully observed but seemingly 
comprised of elongated, clinging, physiognomically unbranched, leafy, non- 
flowering stems and slightly lengthened mostly unbranched, free, leafy, 
flowering stems; stems smooth, flexuous, climbing stems + terete, 
occasionally weakly angled, free stems terete, to similarly sulcate, without 
prophyll. cataphyll and petiolar sheath fibre, internodes 0.3 — 5 x 0.2 — 0.3 
cm on free shoots, flowering shoots with shorter internodes, separated by 



Rhaphidophora in New Guinea, Australia and Western Pacific 



139 



weak straight leaf scars, older stems woody: flagellate foraging stems not 
observed; clasping roots and feeding roots not observed; leaves spiro- 
distichous. slightly scattered; cataphylls and prophylls membranous, 
caducous; petiole grooved adaxially, 1.5 — 5 x 0.1 — 0.18 cm, smooth, with a 
slight apical and basal geniculum; petiolar sheath slightly prominent, 
extending beyond the apical geniculum by two ligules. caducous leaving a 
continuous scar from the petiole base, around the top of the apical 
geniculum and back to the base: lamina entire, elliptic to elliptic-lanceolate 
or oblanceolate, slightly falcate, 5 — 16 x 2.2 — 7 cm. thinly coriaceous, base 
cuneate subovate or minutely cordate, apex acute to briefly acuminate, 
with a small tubule: midrib raised abaxially, sunken adaxially; primary 
venation pinnate, slightly raised on both surfaces in dried material; 
interprimaries reticulate to sub-parallel to, but much less distinctive than, 
primaries, degrading into weakly reticulate venation, very slightly raised 
abaxially; secondary and tertiary venation reticulate; inflorescence solitary, 
subtended by a fully developed foliage leaf and a caducous cataphyll; 
peduncle slightly compressed-terete, 1 — 3 x 0.2 — 0.3 cm; spathe cigar-shaped, 
apex beaked, 1 — 4.5 x 0.7 — 1.2 cm. fleshy, yellow-green to yellow-cream, 
caducous leaving a large scar; spadix cylindrical, sessile, inserted level on 
peduncle, 1.5—2.5 x 0.5 — 0.8 cm. obtuse, white; stylar region rounded- 
rhombohexagonal, 0.9 — 1 x 0.85 — 1.1 mm, truncate: stigma punctiform, c. 
0.3 mm diam.. slightly prominent in dried material: anthers not exserted at 
male anthesis; infructescence not observed. 

Distribution: Papua New Guinea (Bougainville. Manus). Solomon Islands 
(Fauore, Guadalcanal, Kolombangara. Malaita. San Jorge. Small Malaita). 

Habitat: Well-drained primary and secondary forest on flat and hilly terrain, 
occasionally in lowland swamp forest. 3-1000 m altitude. 

Notes: 1 . In herbaria Rhaphidophora mima is much confused with the very 
similar R. neoguineensis (absent from the Solomon Islands) although it is 
readily distinguished by a caducous, cigar-shaped spathe and cylindrical 
spadix (spathe globose and marcescent, spadix globose-ellipsoid in R. 
neoguineensis). 

2. The specific epithet is the adjectival form of mimus, from the Greek. 
mimos, actor, in allusion to the confusing similarity of this species to 
Rh ap hidophora n eogi uneens is . 

3. The Manus collection (Foreman & Katik LAE 59291A), while outside 
the main geographical range of the species, is undoubtedly R. mima. 



140 



Card. Bull. Singapore 53 (2001) 



Other specimens seen: PAPUA NEW GUINEA. Manus Prov.: Manus, near Pelikawa, 
Foreman & Katik LAE 59291A (L, US); North Solomons Prov.: Bougainville, Sulka wide 
bay, Bateson 89, 91, 92, 93 (K); Buin, Kugumaru, Kajewski 1891 (BM, GH, K, SING); 
Koniguri, Kajewski 2162 (GH, P); Pavairi, Lavarack & Lavarack NGF 31091 (K, L, US); 
McKillup's Arawa Plantation, 6 miles west of Kieta, Nicolson 1516 (US), Nicolson 1522 (B, 
K, L, US); vicinity of Aku village, c. 10 miles west of Buin, Schodde 4070 (BRI, CANB, 
GH, K, L, LAE, US); Siwai, Waterhouse 207 (GH, K), Waterhouse 778 (K). SOLOMON 
ISLANDS. Malaita: SW Malaita, Wairokai River area, Gafui et al. BSIP 10215 (BSIP, K, 
SING); Suu area. Mauriasi et al. BSIP 13632 (BSIP. L, SING); Tantalau - Kwalo trail, near 
Kwalo, Stone 2366 (BISH, K, US); Small Malaita: Palasu'u, east of Rota School. Gafui 
BSIP 17296 (BSIP, K, SING); San Jorge: Talise village. Hunt RSS 2722 (K. US); 
Kolombangara: Ridge west of Vila river, Mauriasi et al. BSIP 8429 (BSIP, K, SING): 
Fauore: Halluma River, Mauriasi et al. BSIP 13977 (BSIP, K, L, SING). 

19. Rhaphidophora neoguineensis Engl. 

Rhaphidophora neoguineensis Engl, in K. Schum. & Hollrung, Fl. Kais. 
Wilh. Land (1889) 19 ('neo-guineensis'); K. Schum. & Lauterbach, Fl. 
Schutzgeb. Siidsee (1900) 211; Engl. & K. Krause in Engl., Pflanzenr. 37 
(IV.23B) (1908) 20—21, Fig. 3 ('novo-guineensis'): Engl. & K. Krause, 
Nova Guinea 8 (1912) 805; Alderw., Bull. Jard. Bot. Buitenzorg III, 1 
(1920) 386 ('novo-guineensis"): Alderw., Bull. Jard. Bot. Buitenzorg III, 4 
(1922) 337 — 338 ('novo-guineensis")\ K. Krause & Alderw., Nova Guinea 
14 (1924) 212 ('novo-guineensis'). — Type: Papua New Guinea ('Kaiser 
Wilhelmsland'), Madang Prov., Adelbert Range, near Hatzfeldthafen, Oct. 
1886, Hollrung 372 (B, holo). 

Figure 21 

Slender, leptocaul, homeophyllous liane to 5 m; seedling not observed; pre- 
adult plants forming small terrestrial colonies; adult shoot architecture 
comprised of elongated, clinging, physiognomically unbranched, leafy, non- 
flowering stems and very short to somewhat lengthened mostly unbranched, 
free, sympodial, leafy, flowering stems; stems smooth, flexuous, climbing 
stems ± terete, occasionally weakly sulcate on two opposing sides, free 
stems terete, to similarly sulcate, without prophyll, cataphyll and petiolar 
sheath fibre, internodes 1 — 9 x 0.2 — 0.6 cm on clinging and free shoots, 
flowering shoots with shorter internodes, separated by weak straight leaf 
scars, occasionally disarticulating into 2 — 4 internode lengths and these 
later rooting and forming independent plants, older stems woody; flagellate 
foraging stems frequent, terete in cross-section with reduced leaves basally, 
terminal portion with caducous cataphylls; clasping roots arising sparsely 
from the clinging stems, very slightly pubescent; feeding roots solitary from 



Rhaphidophora in New Guinea, Australia and Western Pacific 



141 




Figure 20. Rhaphidophora mima P.C. Boyce 

A. flowering branch, spathe intact x '/-,: B. flowering branch, spathe fallen x V,:C. inflorescence, 
spathe fallen, x 2: D. spadix detail, post-anthesis x 4. A from Kajewski 1891: B from Foreman 
& Katik LAE 59291 A: CD from Nicolson 1522. 



142 



Card. Bull. Singapore 53 (2001 ) 



nodes, clinging to climbing surface; leaves weakly spiro-distichous, 
moderately densely arranged; cataphylls and prophylls membranous, 
caducous; petiole grooved adaxially, 3 — 12 x 0.1 — 0.2 cm, smooth, with a 
slight apical and basal geniculum, although older leaves often with genicula 
enlarged and cracking-corky; petiolar sheath slightly prominent, extending 
beyond the apical geniculum by two ligules, caducous leaving a continuous 
scar from the petiole base, around the top of the apical geniculum and 
back to the base; lamina entire, elliptic to elliptic-lanceolate or oblanceolate, 
6 — 25 x 2 — 9 cm, thinly coriaceous to submembranous, base cuneate to 
acute or subovate, apex acute to weakly acuminate, with a minute tubule; 
midrib raised abaxially, slightly sunken adaxially in fresh material, slightly 
sunken abaxially and adaxially in dry material; primary venation pinnate, 
slightly raised on both surfaces in dried material; interprimaries reticulate 
to sub-parallel to, but much less distinctive than, primaries, degrading into 
weakly reticulate venation, very slightly raised abaxially; secondary and 
tertiary venation reticulate; inflorescence solitary, subtended by a fully 
developed foliage leaf and a caducous cataphyll; peduncle slightly 
compressed-terete, 1.3 — 2.5 x 0.15 — 0.3 cm; spathe globose to ovoid- 
ellipsoid, truncate basally at insertion on petiole, apex briefly beaked, 1 — 2 
x 1 — 2.5 cm, thick fleshy, dark yellow, marcescent, drying brown, later 
pushed off by developing infructescence leaving a large scar; spadix globose 
to ellipsoid-cylindrical, sessile, inserted level on peduncle, 1 — 1.5 x 0.7 — 1 
cm, obtuse, yellow-white; stylar region rounded-rhombohexagonal, 0.9 — 
1.5 x 0.85 — 1.2 mm, truncate; stigma punctiform to very slightly elliptic, c. 
0.45 x 0.3 mm diam.. prominent in dried material; anthers well-exserted at 
male anthesis; in fructescence not observed. 

Distribution: New Guinea. Widespread but, by the exclusion of many 
collections here recognized as new species, not as common as was once 
thought. 

Habitat: Primary to disturbed secondary lowland, gallery and hill monsoon 
and rain forest on various substrates including coralline limestone. 3-700 
m altitude. 

Note: Rhaphidophora neoguineensis is characterized by a small ( 1 — 2 x 1 — 
2.5 cm), marcescent, globose spathe and ellipsoid-cylindrical spadix. In 
herbaria it is much confused with R. mima but which is readily separated 
by the cigar-shaped, caducous spathe and cylindrical spadix. Confusion 
with R. schlechteri is also possible, although this has a larger spathe (4 — 6 x 
c. 2.4 cm) and a stipitate spadix. 



Rhaphidophora in New Guinea. Australia and Western Pacific 



143 




Figure 21. Rhaphidophora neoguineensis Engl. 

A. adult shoot with flowering branch x "'/.,: B. adult clinging shoot x V 3 : C. flagellate shoot x V 2 : 
D. leaf lamina x V 3 ; E. venation detail x 3: F. inflorescence, spathe removed x 4: G. spadix detail, 
male anthesis x 6. A-C. F-G from Hay K2 [cult. Kew (Acc. no. 1982-5010) & K spirit no. 
58061] x V 3 ; D-E from Nicolson 1402. 



144 



Card. Bull. Singapore 53 (21)01 ) 



Oilier specimens seen: INDONESIAN PAPUA. Bonggo Range, Mamberamo. Pionierbivak. 
Lain 763 (L). Jayapura Prov.: Taritatau (Idenburgh) River, Bernhard Bivak, Meyer Drees 
279 (L); Biak, hill northeast end of Mokmer airport, Nicolson 1565 (L, US). PAPUA NEW 
GUINEA. East Sepik Prov.: Wewak, 2 miles west of But village, Essig & Martin LAE 
55124 (L, LAE): West Sepik Prov.: Vanimo, Wutung, on Papua New Guinea/Indonesian 
Papua border. Streimann LAE 52789 (LAE. US): Sepik Prov.: Sepik River, Gjellerup 330 
(L); Kelel, Schlechter 16329 (P): Keneyia, Schlechter 18298 (P); Western Highlands Prov.: 
Mt Hagan, near Ruti E.L.G., Henty & Streimann NGF 38835 (US): Morobe Prov.: Lae, 
Markham Bridge, Kairo & Streimann NGF 30715 (GH, K, L, SING, US): Lae B.G., Millar 
NGF 9938 (GH, K, L), Nicolson 1402 (B, BM, K. L. P, SING, US); Near Markham River, 9 
miles southwest of Lae on Bulolo road, Nicolson 1480 (US); Central Prov.: Port Moresby. 
Brown River F.R.. 20-25 miles northwest of Port Moresby, Nicolson 1427 (US). 

20. Rhaphidophora okapensis P.C. Boyce & Bogner 

Rhaphidophora okapensis P.C. Boyce & Bogner, Gard. Bull. Singapore 52 
(2000) 94, fig.2. — Type: Papua New Guinea, Eastern Highlands, 5 miles 
NE of Okapa, 24 Sept. 1964, Hartley TGH 13098 (CANB, holo; GH, K, L, 
iso). 

Figure 22 

Moderate, slightly robust, semi-leptocaul. homeophyllous neotenic liane to 
unknown ultimate height; seedling stage and pre-adult plants not observed; 
adult shoot architecture comprised of clinging, physiognomically unbranched, 
densely leafy, sterile stems, and clinging, leafy flowering stems; stems terete 
in cross-section, smooth, without prophyll, cataphyll and petiolar sheath 
remains, internodes to 5 x 0.75 cm, separated by slightly swollen nodes 
with ± sloping scars, older stems sub-woody; flagellate foraging stems not 
observed; clasping roots arising sparsely from the internodes. smooth to 
very slightly pubescent; feeding roots not observed; leaves distichous, those 
on adherent shoots weakly shingling to slightly scattered, those on free 
shoots pendent to slightly spreading; cataphylls and prophylls membranous, 
caducous; petiole shallowly and broadly grooved, 1 — 4 x 0.1 — 0.2 cm, 
smooth, apical and basal genicula quite prominent; petiolar sheath 
prominent, membranous, ligulate, ligule extending c. 1.5 cm beyond apical 
geniculum, soon drying and adhering to stem, later disintegrating and falling; 
lamina ovate, adaxially mid-green, abaxially paler, very stiffly coriaceous, 
2.5 — 8.5 x 0.9 — 4 cm, base cordate to rounded and slightly notched, apex 
long acuminate with a pronounced tubule; midrib proximally raised 
abaxially. slightly impressed adaxially; primary venation pinnate, slightly 
raised on both leaf surfaces; interprimaries sub-parallel to and barely 
distinguishable from primaries, slightly raised on both leaf surfaces; 
secondary venation reticulate, slightly raised abaxially and adaxially; 



Rhaphidophora in New Guinea, Australia and Western Pacific 



145 




Figure 22. Rhaphidophora okapensis P.C. Boyce & J. Bogner 

A. adult shoot with flowering branch x V 4 ; B. leaf lamina x 1: C. venation detail x 4; D. 
inflorescence x l'/ : : E. spadix detail at male anthesis x 10: F. pistil, side view x 10. All from 
Hartley 13098. 



146 



Card. Bull. Singapore 53 12001) 



in florescence solitary on short leafy shoots, subtended by a fully developed 
or reduced foliage leaf, and a soon-degrading, membranous, long-ligulate 
prophyll; peduncle terete, 4 — 6 x 0.15—0.2 cm; spathe broadly canoe-shaped, 
rounded, minutely apiculate, 3 — 3.5 x 1.2 — 1.5 cm, stiffly fleshy, yellow, 
turning purple, (early marcescent?) leaving a large scar at the base of the 
spadix; spadix stout, cigar-shaped, stipitate, inserted level on stipe, 1.9 — 
2.2 x 0.75 — 1 cm; stipe 4 — 4.5 x 1 — 1.2 mm; stylar region conical, regularly 
rhombohexagonal in plan view, 0.8 — 1 x c. 1 mm; stigma slightly raised, 
punctiform, c. 0.4 mm diam.; anthers exserted at male anthesis; infructescence 
not seen. 

Distribution: Papua New Guinea (Eastern Highlands and Central 
Provinces). 

Habitat: Disturbed mixed forest on slope. 1200-1600 m altitude. 

Note: A very distinctive species notable for the stiffly erect, densely leafy 
stems, the thickly coriaceous, ovate-cordate leaves with a long acuminate 
tip, the relatively large inflorescences turning purple at maturity, the long- 
stipitate spadix, and the conical stigmatic region. 

Other specimens seen: PAPUA NEW GUINEA. Central Prov.: Boridi. Can 14863 (BM, L. 
SING); Eastern Highlands Prov.: Kainantu, Arau-Andandara road, Slreiman NGF 23963 
(US). 

21. Rhaphidophora pachyphylla K. Krause 

Rhaphidophora pachyphylla K. Krause, Bot. Jahrb. Syst. 49 (1912) 92; P.C. 
Boyce & Bogner, Gard. Bull. Singapore 52 (2000) 94, fig.2. — Type: Papua 
New Guinea, Madang Prov., near Wabbe, 29 Aug. 1907, Schlechter 16463 
(B, holo; P, iso). 

Figure 23 

Moderate to rather large, somewhat robust, semi-leptocaul, homeophyllous 
neotenic liane to 5 m; seedling stage a non-skototropic shingling juvenile 
shoot; pre-adult plants forming small terrestrial colonies; adult shoot 
architecture comprised of clinging, physiognomically unbranched, mostly 
densely leafy, sterile stems and free, leafy flowering stems; stems rectangular- 
terete in cross-section, widest side prominently convex, smooth, mid-green, 
without prophyll, cataphyll and petiolar sheath fibre although flowering 
shoots occasionally with parchment-like remains, internodes to 3 x 1 cm, 



Rhaphidophora in Sew Guinea, Australia and Western Pacific 



147 




Figure 23. Rhaphidophora pachyptnlla K. krause 

A. adult shoot with flowering branch x '7 : : B. section of adult climbing shoot x V : : leaf lamina 
x V : : C. juvenile climbing shoot x V,; D. pre-adult climbing shoot x 7 4 : E. leaf lamina x 3 / 4 : F. 
venation detail x 4: G. inflorescence, spathe fallen x 1 'A: H. spadix detail, post-male anthesis x 
10. A. E - F from Krause 108: B from Kalkman 3389: C-D from Nicolson 1416: G-H from 
Schlechter 16436. 



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Gard. Hull. Singapore 53 12001) 



separated by rather prominent + straight scars, older stems sub-woody; 
flagellate foraging stems absent; clasping roots arising sparsely from the 
internodes, pubescent; feeding roots not observed; leaves distichous, those 
on adherent shoots shingling, those on free shoots slightly spreading, all 
densely arranged; cataphylls and prophylls membranous, caducous; petiole 
deeply grooved, 1.5 — 4 x 0.15 — 0.2 cm, smooth, apical and basal genicula 
quite prominent; petiolar sheath prominent, membranous, ligulate, slightly 
unequal on one side, of short-duration, degrading to very weak fibres and 
soon falling; lamina narrowly ovate-elliptic, stiffly coriaceous, base acute 
to cuneate, apex acute with a short tubule; midrib prominently raised 
abaxially, slightly impressed adaxially; primary venation pinnate, slightly 
raised abaxially, somewhat impressed adaxially; interprimaries sub-parallel 
to primaries, slightly raised on both leaf surfaces; secondary venation weakly 
reticulate, slightly raised abaxially, + flush adaxially; inflorescence solitary 
on short to somewhat elongated leafy shoots, subtended by a fully developed 
foliage leaf, and soon-degrading membranous prophyll; peduncle terete, 

4 — 7 x 0.2 — 0.25 cm; spathe broadly canoe-shaped, obtuse to slightly pointed, 

5 — 6.5 x 1.5 — 2.2 cm, stiffly fleshy, yellow, gaping at female anthesis and 
then caducous leaving a large scar at the base of the spadix; spadix narrowly 
cigar-shaped, long stipitate, inserted obliquely on stipe, 4 — 5.5 x 1.2 — 1.6 
cm, yellow; stipe 6 — 10 x 2 — 2.4 mm; stylar region weakly developed, mostly 
irregularly rhombohexagonal, 1 — 1.2 xl — 1.4 mm, truncate; stigma slightly 
raised, punctiform, c. 0.2 — 0.3 x 0.3 — 0.35 mm; anthers slightly exserted at 
male anthesis; infructescence not seen. 

Distribution: Indonesian Papua, Papua New Guinea. The scattered localities 
and few collections suggest a widespread but uncommon species. 

Habitat: Lowland monsoon or rain forest at 10-30 m altitude. 

Notes: 1. Long confused with Rhaphidophora hayi, R. pachyphylla differs 
in leaf shape, and form of the stigma. Additionally, R. pachyphylla lacks 
the disarticulating shoots unique to R. hayi. and never has foraging shoots. 

2. It is still not fully clear whether R. pachyphylla as here defined is a 
single taxon. Hoogland & Schodde 6943 (Western Highlands Prov., Wabag, 
near Poio village, west slopes of lower Yaki valley, 6 July 1960, in montane 
Nothofagus forest, 2380 m (BM, GH, L), while vegetatively matching typical 
R. pachyphylla, has a markedly rounded spathe apex, a shorter stipe (c. 3 
mm long) and also represents an enormous altitudinal increase on the 
other specimens seen. More specimens of "R. pachyphylla' from higher 
altitude are required to resolve this plant's status. 



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Other specimens seen: INDONESIAN PAPUA. Tamimonding. Kalkman s.n. (L); Digul 
Prov.: Merauke. Bis, Agats. Widjaja 6344 (BO, K. L). PAPUA NEW GUINEA. Central 
Prov.: Brown River F.R.. 20 - 25 miles northwest of Port Moresby. Nicolson 1416 (K, L. 
US). 

22. Rhaphidophora petrieana A. Hay 

Rhaphidophora petrieana A. Hay. Telopea 5 (1993) 295. fig. 1. — Type: 
Australia. Queensland, National Park Reserve 904, Palmerston Highway, 
30 Nov. 1982, B. Gray 2862 (QRS. holo). 

Rhaphidophora sp. aff. australasica (Qld) in Jones & Gray. Climbing PL 
Australia (1988) 316, unnumbered plate p. 322. 

Figure 24 

Medium to large, moderately robust, semi-leptocaul homeophyllous liane 
to 20 m; seedling and pre-adult plants not observed: adult shoot architecture 
comprised of greatly elongated, clinging, physiognomically unbranched. 
sparsely leafy, non-flowering stems and long, moderately elaborated, free, 
sympodial, densely leafy, flowering stems: stems smooth, climbing and free 
stems terete in cross-section, green, later mid-brown, without prophyll. 
cataphyll and petiolar sheath fibre, internodes to 2.5 — 5 x 0.5 — 1 cm on 
clinging shoots, usually less stout on free shoots, separated by well defined, 
slightly oblique, corky leaf scars, older stems subwoody; flagellate foraging 
stems absent: clasping roots sparse, arising from the nodes of clinging stems, 
minutely pubescent: feeding roots not observed; leaves spiro-distichous on 
clinging shoots, disticho-secund on free shoots: cataphylls and prophylls 
membranous, very quickly drying and falling: petiole grooved adaxially. 
4.5 — 8 x 0.15 — 0.3 cm, smooth, apical and basal genicula weakly defined; 
petiolar sheath very prominent, extending to just below apical geniculum. 
swiftly drying and eventually falling more-or-less entire leaving a 
conspicuous scar; lamina entire, narrowly ovate to lanceolate, somewhat 
falcate, slightly oblique, 4.5 — 22.5 x 1 — 4.5 cm. thinly coriaceous, base acute, 
apex acuminate, with a tiny tubule: midrib raised abaxially. slightly sunken 
adaxially; primary venation pinnate, slightly raised abaxially and adaxially; 
interprimaries parallel to primaries and barely less prominent, very slightly 
raised abaxially and adaxially; secondary venation ± obscure in fresh 
material, visible as a faint reticulum in dried specimens; inflorescence 
solitary, subtended by a partially developed foliage leaf and a membranous 
cataphyll; peduncle compressed-terete, 3 — 10 x 0.15 — 0.8 cm; spathe canoe- 
shaped, stoutly short- to rather long-beaked, 6 — 8 x 1 — 3 cm (8 cm wide 
flattened out), thickly stiff-fleshy, creamy yellow, caducous at female 



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Card. Bull. Singapore 53 (2001) 




( 



Figure 24. Rhaphidophora petrieana A. Hay 

A. flowering shoot x 2 / 5 ; B. inflorescence, longitudinal section x 7,; C. spadix detail, male anthesis 
x 6. All from Hyland 9162. 



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anthesis: spadix bluntly tapering-cylindric. stipitate. inserted + level on 
stipe. 3 — 6 x 1 — 2.5 cm; stipe terete, c. 1 x 0.6 cm; stylar region mostly 
rounded-hexagonal, c. 2 x 1.8 — 2 mm. conical: stigma punctiform and 
prominently raised, c. 0.25 — 0.3 mm diam.; anthers exserted at male anthesis: 
infructescence not observed. 

Distribution: Australia (wet tropical regions of eastern Queensland). 

Habitat: Lowland to lower montane rain forest. Sea level to 800 m altitude. 

Note: Long confused in herbaria and literature with Rhaphidophora 
australasica but readily distinguished by flowering on free lateral shoots 
and by the stipitate spadix. Further, in R. petrieana the petiolar sheaths fall 
more-or-less intact and do not degrade into semi-persistent fibres as they 
do in R. australasica. 

Other specimens seen: AUSTRALIA. Queensland: Foot of Mt Demi. Flecker 9005 (QRS): 
Timber Reserve 55. Whyanbeel. Gray 202 (QRS): State Forest Reserve 310, Upper 
Goldsborough Logging Area. Gray 1055 (QRS): Cooper Creek, Hind 246 (NSW); State 
Forest Reserve 755, Barong Logging Area, Hyland 9162 (BRI. L, NSW. QRS). 

23. Rhaphidophora pilosa P.C. Boyce. sp. nov. 

Rhaphidophora pilosa R. gorokensi R. kokodcnsicjuc simillima est, ab ilia 
apices surculorum fibrosos (fibram in R. gorokensi deest) habenti atque ad 
hac natura coacta (non simpliciter fibrosa) reliquiarum apicum surculorum 
differens. Cum R. waria earn confundere potest, quamquam ilia species 
magis major est in surculis adhaerentibus non liberis florens. — TYPUS: 
Indonesian Papua, Kepala Burung Prov., Arfak Mts, Minjambau. 20 May 
1962, C. Versteegh BW 12647 (L, holo: MAN. iso). 

Figure 25 

Small, homeophyllous (?) liane to unknown ultimate height; seedling stage 
and pre-adult plants not observed; adult shoot architecture comprised of 
elongated, clinging, physiognomically unbranched, leafy(?), non-flowering 
stems and short, free, sympodial, flowering stems; stems terete in cross- 
section, internodes to at least 5 x 0.5 cm, separated by straight oblique, 
weak leaf scars; flagellate foraging stems, unknown; clasping roots solitary, 
stout, arising from nodes; feeding roots not observed; leaves weakly spiro- 
distichous on free shoots; cataphylls and prophylls chartaceous, very soon 
degrading into weak fibres and patches of soft felt-like debris; petiole weakly 
canaliculate, 4.5 — 9 x 0.2 — 0.25 cm. smooth, apical and basal geniculum 




Figure 25. Rhaphidophora pilosa P.C. Boyce 

A. flowering branch, spathe intact x : /c B. flowering branch, spathe fallen x 2 / 5 ; C. spadix detail, 
post-anthesis x 5. All from Versteegh BW 12647. 



Rhaphidophora in Mew Guinea, Australia and Western Pacific 



153 



well defined petiolar sheath prominent, chartaceous. extending to apical 
geniculum, very swiftly degrading to weak fibres and strips of soft felt-like 
debris: lamina entire, narrowly lanceolate to narrowly lanceolate-elliptic, 
occasionally very slightly falcate. 6 — 18 x 2 — 3.5 cm. stiffly coriaceous, 
base rounded to acute, apex long-acuminate with a prominent apicule: 
midrib raised abaxially. slightly flush to very slightly sunken adaxially: 
primary venation densely pinnate, very slightly raised abaxially and 
adaxially: interprimaries parallel to primaries, slightly less pronounced: 
secondary venation very faintly open-reticulate: inflorescence solitary, 
subtended by a foliage leaf and felted debris: peduncle stout, compressed- 
terete. 8 — 11 x 0.3 — 0.5 cm; spathe canoe-shaped, very long stout-acuminate. 
9.5 x c. 1.8 cm. marcescent: spadix cigar-shaped, sessile, 5.5 x 1 cm; stylar 
region rhombohexagonal. c. 0.9 — 1 x 1.2 — 1.5 mm. truncate: stigma 
punctiform. flush, c. 0.4 mm diam.: anthers not exserted at male anthesis: 
infructescence cylindrical, c. 8 x 1.8 cm. 

Distribution: Indonesian Papua (Kepala Burung Prov.). Known only from 
the type. 

Habitat: Secondary submontane forest. 1250 m altitude. 

Notes: 1. Rhaphidophora pilosa is one of several montane species with 
stiffly coriaceous leaves (Okapensis Group). It is most similar to R. 
gorokensis and R. kokodensis, differing from the former in possessing 
fibrous shoot tips (R. gorokensis lacks fibre) and from the latter in the 
felted (not simply fibrous) nature of the shoot tip debris. Confusion with 
R. waria is possible although this is a much larger species flowering on 
clinging, not free, shoots. 

2. The specific epithet is from the Greek 'pilosa\ felt-like, in allusion to 
the unique manner in which the cataphylls. prophylls and petiolar sheath 
degrade into felted fibres. 

24. Rhaphidophora schlechteri K. Krause 

Rhaphidophora schlechteri K. Krause, Bot. Jahrb. Syst. 49 (1912) 94. — 
Type: Papua New Guinea. Eastern Highlands Prov.. Tanqueti camp, 27 
Nov. 1908. Schlechter 18888 (B. holo). 

Rhaphidophora conferta K. Krause, Bot. Jahrb. Syst. 49 (1912) 95, synon. 
nov. — Type: Papua New Guinea. Keneyia Camp. 2 Oct. 1908. Schlechter 
18308 (B.'holo: P, iso). 



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Gard. Bull. Singapore 53 (2001) 



Rhaphidophora peekelii Engl. & K. Krause, Bot. Jahrb. Syst. 54 (1916) 78; 
Peekel ex Henty, Fl. Bismarck Archipelago (1984) 66 — 67 fig. 110, synon. 
nov. — Type: Papua New Guinea, New Ireland, Namatanai, Peekel 296 (B, 
holo). 

Rhaphidophora buergersii Engl. & K. Krause, Bot. Jahrb. Syst. 54 (1916) 
78 { l biirgersu), synon. nov. — Type: Papua New Guinea, Sepik Prov., 
Lordberg, 9 Dec. 1912, Ledermann 10241 (B, holo). 

Rhaphidophora drepanophylla Alderw., Bull. Jard. Bot. Buitenzorg III, 4 
(1922) 340; K. Krause & Alderw., Nova Guinea 14 (1924) 213, synon. nov. 
— Type: Indonesian Papua, Bonggo Range, Mamberamo, near Prauwen 
Bivak, 1 Sept. 1920, Lam 1014 (BO, holo; L, iso). 

Rhaphidophora obliquata Alderw., Bull. Jard. Bot. Buitenzorg III, 4 (1922) 
340; K. Krause & Alderw., Nova Guinea 14 (1924) 212, synon. nov. — 
Type: Indonesian Papua, Bonggo Range, Mamberamo, near Pionier Bivak, 
30 June 1920, Lam 483 (BO, holo). 

Figures 26 & 27 

Slender to moderately robust, leptocaul, homeophyllous (?) liane to 20 m; 
seedling and pre-adult plants not observed; adult shoot architecture comprised 
of elongated, clinging, physiognomically unbranched, leafy, non-flowering 
stems and very short to somewhat lengthened, unbranched, free, sympodial, 
leafy, flowering stems; stems smooth, flexuous, terete, without prophyll, 
cataphyll and petiolar sheath fibre, internodes 1 — 9 x 0.5 — 1.5 cm, separated 
by thin, slightly oblique, leaf scars, older stems woody; flagellate foraging 
stems absent; clasping roots arising singly from the node; feeding roots not 
observed; leaves spiro-distichous, scattered; cataphylls and prophylls 
membranous, caducous; petiole grooved adaxially, 7 — 14 x 0.2 — 0.3 cm, 
smooth, with a very slight apical and prominent basal geniculum; petiolar 
sheath very prominent on newest leaves, very broad basally, narrowing c. + 
way along petiole and extending beyond the apical geniculum by two tiny 
ligules, caducous leaving a continuous scar from the petiole base, around 
the top of the apical geniculum and back to the base; lamina entire, 
lanceolate to oblong-lanceolate, slightly to prominently oblique, 10 — 34 x 
4 — 9.5 cm, thinly coriaceous, base obtuse-cuneate to rounded and minutely 
cordate, apex falcate, acute to long-acuminate, with a tiny tubule; midrib 
slightly raised abaxially, flush to very slightly sunken adaxially; primary 
venation pinnate, slightly raised on both surfaces; interprimaries sub-parallel 
and barely less distinctive than primaries; secondary and tertiary venation 
reticulate, conspicuous in dry material; inflorescence solitary, subtended by 



Rhaphidophora in New Guinea, Australia and Western Pacific 



155 




Figure 26. Rhaphidophora schiechteri K. Krause 

A. flowering shoot x V,; B. leaf lamina x V,; C. venation detail x 4; D. inflorescence, spathe 
fallen x 1; E. spadix detail, pre-anthesis x 8. A & C from Schlechter 18888: B. D-E from Boraule 
et al. BSIP 9141. 



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Card. Bull Singapore 53 (2001) 



a incompletely developed foliage leaf and one or more prominent caducous 
cataphylls; peduncle slightly compressed-terete, 5 — 9 x c 0.5 — 1 cm; spathe 
cigar-shaped, apex stoutly beaked, 4 — 11 x c. 3.5 cm (flattened out), thick, 
marcescent, falling as fruits ripen, green at male anthesis; spadix cylindrical, 
stipitate, inserted slightly obliquely on stipe, 6 — 8.5 x 1.5 — 2 cm, creamy 
white; stipe terete, 0.5 — 1 cm long; stylar region depressed- 
rhombohexagonal, c. 1.2 x 1.3 mm, truncate; stigma punctiform, c. 0.3 mm 
diam., raised at male anthesis but barely prominent in dried material; 
anthers strongly exserted at male anthesis; infructescence stoutly cylindrical, 
c. 8 x 3 cm, with dry spathe persisting. 

Distribution: Widespread from Papua New Guinea (including Bismarck 
Archipelago) to the Solomon Islands. 

Habitat: Primary to disturbed secondary broadleaf and coniferous 
(Araucaria) forest on humus, alluvium and lava. Sea level to 1500 m altitude 

Notes: 1. As defined here Rhaphidophora schlechteri is a variable and 
widespread species. The broad species concept adopted is based on 
examining all the numerous duplicates of the Nicolson collections cited 
here that exhibit wide variation, which supports the merging of elements 
recognized as separate species by earlier workers. 

2. In most of its medium to large manifestations R. schlechteri is similar in 
overall appearance to R. mima, but is distinguishable by the marcescent 
spathe and stipitate spadix. Small forms of R. schlechteri maybe confused 
with R. neoguineensis, although the globose spathe and sessile ellipsoid 
spadix readily distinguish the latter. 

Other specimens seen: PAPUA NEW GUINEA. West Sepik Prov.: Telefomin, Sanduan, 
track on leaf bank of Mai (Yuwa) river, to 0.5 km south of Fiak airstrip, Frodin el at. 2553 
(K); Central Prov.: Sogeri Plateau. Rouna Waterfall, 20 miles east of Port Moresby, Nicolson 
1432 (B, BM, K. L. P. SING, US); Morobe Prov.: Wau, road half way to Yamap, Kairo 
NGF 44078 (L, NGF, US); 10 miles north of Bulolo. Nicolson 1477 (B. K, L, P. US): 
Madang Prov., southern slopes of Finisterre Range, near Budemu, Pullen 5996 (BM, L. 
LAE); Saidor, Matafuma village, vicinity of Pukie Primary School, Vandenberg & Katik 
NGF 42372 (GH, K, L. US); North Solomons Prov.: Bougainville, vicinity of Barilo village, 
c. 6 miles north of Buin Station, Schodde 3935 (GH, L); New Ireland: Mussau, K0ie & 
Olsen 1377 (L); Namatanai, coastal region, near Tamul River, c. 0.5km NW of Taron, east 
coast, Sands et al. 2044 (GH, K, L, US); SOLOMON ISLANDS. Guadalcanal: West 
Guadalcanal, Wanderer Bay. Vuragoba area. Boraule et al. BSIP 9141 (BSIP, K, SING); 
North central Guadalcanal, Tina River, Nakisi & Babala BSIP 8222 (BSIP, K, SING); San 
Cristobal: Puepue River, Brass, 2794 (BO, GH, L); South Ridge west of Tetere village, 
Gafui et al. BSIP 1 2575 (BSIP. K. SING); Fauore ("Fauo - ) Island: Guppy 196 (K); Ulawa: 
Moli, Teona BSIP 6303 (BSIP, K. L. SING). 



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25. Rhaphidophora spathacea Schott 

Rhaphidophora spathacea Schott, Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 129. 

— Type: 'New Guinea', Zippelius s.n. (L, holo). 

Rhaphidophora apicidata K. Krause, Bot. Jahrb. Syst. 49 (1912) 93, synon. 
nov. — Type: Papua New Guinea ('Kaiser Wilhelmsland'), Madang Prov., 
Finisterre Range, 2 Sept. 1908, Schlechter 18152 (B, holo; P, iso). 

Rhaphidophora palauensis Koidz., Bot. Mag. Tokyo 30 (1916) 400. synon. 
nov. — Type: Palau, Angaur Island, Kayima 154 (TI, holo). 

Rhaphidophora forbesii Rendle, J. Bot. 61, Suppl. (1923) 58, synon. nov. 

— Type: Papua New Guinea, Sogeri Prov., 1885-1886, Forbes 323 (BM, 
holo). 

Rhaphidophora engleri Kanehira, Fl. Micrones. (1933) 409. — 
Rhaphidophora palauensis Engl. & K. Krause, Bot. Jahrb. Syst. 56 (1921) 
433, nom. illeg., non Koidz. (1916). synon. nov. — Type: Palau, Babelthuap 
('Babelthaob'). Dorfe Ngarsul, 21 Feb. 1914, Ledermann 14336 (B, holo). 

Rhaphidophora kanehirae Hatusima. J. Japanese Bot. 15 (1939) 19, fig. 1, 
g — j, synon. nov. — Type: Federated States of Micronesia, Yap. Kanehira 
1188 (TI, holo). 

Figures 28 & 29 

Moderately robust, medium-sized pachycaul, homeophyllous liane to 15 
m; seedling stage not observed; pre-adult plants forming scattered terrestrial 
colonies; adult shoot architecture comprised of clinging, physiognomically 
unbranched, densely leafy flowering stems; stems smooth, mid-green, with 
cataphylls and prophylls persistent and drying dark yellow, then degrading 
into fibres and parchment-like remains, internodes 1- — 6 x 0.4 — 1.5 cm, 
separated by prominent slightly oblique leaf scars; flagellate foraging stem 
absent; clasping roots densely arising from the nodes and internodes, smooth 
and drying with parchment-like epidermis; feeding roots not observed; leaves 
spiro-distichous to distichous; cataphylls and prophylls chartaceous, later 
degrading into strips of tissue and fibres, especially at tips of flowering 
shoots; petiole deeply canaliculate, 8 — 53 x OA — 0.8 cm, smooth with faint 
to rather prominent dark dense speckling, apical geniculum prominent, 
basal geniculum slightly less so; petiolar sheath extending to apical 
geniculum, broad, chartaceous, short-persistent, degrading to sparse fibres 
and papery strips, then falling; lamina entire, ovate-elliptic to oblong- 



Rhaphidophora in Sew Guinea. Australia and Western Pacific 



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Figure 28. Rhaphidophora ^pathacea Schott 

A. flowering shoot x V a : B. leaf lamina x ] / : .; C. venation detail x 3: D. inflorescence, spathe 
removed x V^j E. spadix detail, post-male anthesis x 8: F. stylar region and stigma, side view x 
6. All from Forbes 323. 



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Card. Bull. Singapore 53 (2001) 



lanceolate or oblong-elliptic, slightly oblique, 11 — 35 x 2.5 — 18 cm, 
chartaceous to coriaceous, base rounded to acute, apex acute to acuminate; 
midrib prominently raised abaxially, + flush adaxially; primary venation 
pinnate, raised abaxially, less so adaxially; interprimaries sub-parallel to 
primaries, slightly to much less prominent than the primaries, slightly raised 
abaxially, barely visible adaxially; secondary venation reticulate, raised, 
especially conspicuous in dry material; tertiary venation minutely reticulate, 
hardly visible; inflorescences two, three or more together, each subtended 
by a prominent chartaceous prophyll and one or more chartaceous 
cataphylls, the entire synflorescence emerging from a mass of dried, 
chartaceous cataphyll remains; peduncle slender to stout, terete, partially 
to completely obscured by cataphylls, 4 — 12 x 0.2 — 0.8 cm; spathe slender 
canoe-shaped, hardly to stoutly beaked, 5 — 13.5 x 2 — 2.5 cm, stiff-fleshy, 
lower 0.7 — 1 cm with connate margins {fide Nicolson 1580. but not observed 
elsewhere), yellow to yellow green, marcescent to early fruiting, eventually 
falling leaving a prominent scar; spadix stoutly cylindrical, sessile, inserted 
almost level on peduncle, 3 — 16.5 x 1 — 1.5 cm, white at male anthesis; 
stylar region conical, mostly hexagonal in top view, 0.9 — 1.2 x 1 — 1.1 mm; 
stigma punctiform, very prominently raised, those at the tip of the spadix 
even more raised, 0.1 — 0.2 x c. 0.3 mm, glossy and almost black in dried 
material; anthers exserted at male anthesis; infructescence stoutly oblong- 
cylindrical, 6 — 9 x 1.4 — 2 cm. 

Distribution: Palau, Indonesian Papua, Papua New Guinea (including 
Woodlark (Muyua) Island), Federated States of Micronesia (Yap). 

Habitat: Lowland to lower montane primary to disturbed secondary forest 
on a variety of substrates including granite, coralline limestone, volcanically- 
derived soils. Sea-level to 1500 m altitude. 

Notes: 1. Most similar to Rhaphidophora versteegii but readily separated 
by the non-shingling juvenile phase and on drying the strongly discolorous 
leaf laminas of which the abaxial surface is bright orange-brown with 
copious, minute tannin cells. Further, R. spathacea never has perforated 
leaf laminas. 

2. In publishing R. spathacea, Schott stated that it was from Java, although 
both type sheets in L are labelled 'Nov. Guinea.' The types of R. spathacea 
match very well collections from New Guinea made under the various 
synonyms proposed above. 



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161 




Figure 29. Rhaphidophora spathacea Schott 

A. flowering shoot x l / 3 ; B. leaf lamina x '/,; C. venation detail x 2; D. pre-adult clinging shoot x 
V 4 ; E. inflorescence, spathe sectioned x 1; F. spadix detail, post male anthesis x 8; G. stylar 
regions and stigmas at spadix tip x 8. All from Brass 23847. 



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Card. Bull Singapore 53 [2001) 



3. The type of R. engleri has rather narrow leaves, although other collections 
from Palau match collections of R. spathacea from New Guinea. 

4. The holotype of R. apiculata K. Krause is notable for the ovate-elliptic 
leaf laminas with primary and interprimary veins almost indistinguishable. 
However, the isotype (P) is of a much longer, narrower leaf, which is much 
closer in appearance to the type of, for example, R. versteegii and R. 
spathacea. 

5. The type of R. forbesii is very incomplete. Nonetheless, the leaf lamina 
shape and spadix match the type of R. apiculata almost exactly. 

Other specimens seen: PALAU: Babelthuap: trail along Japanese pipeline between Ngardmau 
and waterfront, Bowden-Kerby 5983 (US); Lake Ngardok Fosberg 32572 (US); Aulupse'el, 
Dii'ebachal Beach, Evans 581 (US); Yap: Talgo n Bibau village agroforest, Falanrum & 
Faimaw 6795 (US): Angaur: East coast. Fosberg 31986 (US): Angaur, Koidzumi s.n. (TI). 
INDONESIAN PAPUA. Kepala Burung Prov.: Surroundings of Ayawasi, Ave 4736 (BO. 
L); Triton Bay. Le Guillou 'Triton 48' (P); 2 km north of Manokwari, west end of Tafelberg 
F.R., Nicolson 1580 (B. K. L. P. US): Mimika Prov.: Freeport Concession Area, path east at 
Mile 50 on road to Tembagapura, Coode 8037 (BO, K, MAN); Golf Course surrounds. 
Johns 9967 (BO, K, MAN); Freeport Concession Area, Johns 10393 (BO, K, MAN); Golf 
Course surrounds. Johns 10431 (BO. K, MAN). PAPUA NEW GUINEA. East Sepik 
Prov.: vicinity of Malu and April river, Ledermann 7384 (B spirit); Western Prov.. Palmer 
River, 2 miles below junction of Black River. Brass 7100 (GH), Brass 7380 (GH); Central 
Prov.: Port Moresby, above Boridi village. Foreman & Vinas LAE 60099 (BRI, L, LAE. 
US): Morobe Prov.: Along road to Sankwep SE of Lae, c. 10 km beyond Sankwep, Croat 
52808 (MO); North slopes of Mt Missim (Misson). Croat 52952 (K. MO): Bumbu Logging 
Area. 7 miles north of Lae, Nicolson 1386 (B, P. SING. US). Nicolson 1409 (SING, US); 
Lae, Sankwep. 10 miles east of Lae. Streimann NGF 47654 (US); Milne Bay Prov.: Bibiguni 
Camp, Gwariu River, Brass 23847 (GH, L, US); Alotau, Kulumadau Island, Kairo 237 
(GH. L): Woodlark (Muyua) Island: Kulumadau, Brass 28618 (L). 

26. Rhaphidophora spuria (Schott) Nicolson 

Rhaphidophora spuria (Schott) Nicolson, Allertonia 1 (1978) 348. — 
Cuscuaria spuria Seem, ex Schott, Bonplandia 9: 260, nom. nud.\ Schott, 
Bonplandia 9 (1861) 367; Seeman, Viti (1862) 444 & Fl. Vit. (1868) 287; 
Engl., DC, Monogr. Phan. 2 (1879) 251, in syn. pro Cuscuaria maranti folia 
Schott; Engl. & K. Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 68, in 
syn. pro Scindapsus cuscuaria (Aubl.) Presl. — Type: Fiji, Viti Levu, 1860, 
Seeman 655 (K, holo). 

Rhaphidophora storckiana Schott, Bonplandia 10 (1862) 346; Seeman, Fl. 
Vit. (1868) 287; Engl. & K. Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 



Rhaphidophora in New Guinea, Australia and Western Pacific 



163 



43; Parham. PI. Fiji Isl. (1964) 267: ed.2 (1972) 363. — Rhaphidophora 
peepla var. storckiana (Schott) Engl.. DC. Monogr. Phan. 2 (1879) 243; 
Drake. 111. Fl. Ins. Mar. Pac. (1892) 326. — Type: Fiji, Ovalau, 1862. Storck 
911 (K, holo; BM iso). 

Rhaphidophora reineckei Engl.. Bot. Jahrb. Syst. 25 (1898) 9: Engl. & K. 
Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 41. synon. nov. — Type: 
Western Samoa. Savai'i. 1894. Reinecke 594 (B. holo). 

Rhaphidophora graeffei Engl.. Bot. Jahrb. Syst. 25 (1898) 9: Engl. & K. 
Krause in Engl.. Pflanzenr. 37 (IV.23B) (1908) 40, synon. nov. — Type: 
Western Samoa. Upolu. Graeff73 (B. holo). 

Figure 30 

Moderate to large, robust, semi-pachycaul homeophyllous liane to 15 m: 
seedling stage not observed; pre-aduh plants forming small terrestrial 
colonies: adult shoot architecture comprised of elongated, clinging, 
physiognomically unbranched. leafy, non-flowering stems and short, usually 
unbranched. free, sympodial. densely leafy, flowering stems: stems smooth, 
terete in cross-section, with sparse prophyll. cataphyll and petiolar sheath 
fibre, this soon falling, internodes 0.5 — 8 x 0.5 — 2 cm on clinging shoots, 
usually shorter and stouter on free shoots, separated by large, straight, 
corky leaf scars, older stems woody: flagellate foraging stems absent: clasping 
roots densely arising from the nodes and internodes of clinging stems, 
notably pubescent: feeding roots not observed: leaves weakly spiro-distichous 
on clinging and free shoots: cataphylls and prophylls membranous, quickly 
drying and degrading into sparse fibres, these soon falling: petiole deeply 
canaliculate. 14 — 66 x 0.3 — 0.1.5 cm. apical and basal geniculum moderately 
prominent: petiolar sheath prominent, extending to the apical geniculum. 
swiftly drying and degrading into sparse, soon-falling fibres: lamina entire, 
ovate-oblong to oblong-lanceolate or oblong-elliptic, sometimes slightly 
oblique, occasionally falcate. 5.7 — 76 x 2.5 — 32 cm. thinly to quite coriaceous 
(larger laminas tending to be thinner textured), often drying strongly 
discolorous. adaxially mid-brown, abaxially pale brown, base unequal, 
cuneate to rounded, subtruncate or weakly cordate, where present basal 
'lobes" more developed on one side, apex acute to obtuse, acuminate with 
a prominent, short tubule: midrib prominently raised abaxially, ± sunken 
adaxially; primary venation pinnate, raised abaxially and adaxially; 
interprimaries sub-parallel to primaries, hardly less prominent, slightly raised 
abaxially and adaxially: secondary venation reticulate, slightly raised; 
in florescence solitary to several together, strongly sweet-fragrant, if solitary 



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Cant. Bull. Singapore 53 12001) 



then subtended by a partially to fully developed foliage leaf, if more then 
one than subsequent inflorescences each subtended by a soon-degrading 
membranous prophyll and cataphyll; peduncle compressed-terete, often 
with a deep longitudinal sulcus on the shoot side, 5 — 19 x 0.25 — 1 cm; 
spathe broadly canoe-shaped, stoutly beaked, 9.5 — 16 x 2.5 — 5.5 cm (up to 
c. 7 cm wide when flattened out), stiff-fleshy, cream at male anthesis, 
caducous leaving a large, straight scar; spadix cylindrical, inserted ± level 
to somewhat obliquely on peduncle, 6.2 — 15.5 x 1.2 — 1.5 cm, creamy white 
at male anthesis; stylar region mostly hexagonal, 1.6 — 2.4 x c. 2 mm diam., 
truncate, area around stigma sunken; stigma punctiform to slightly ellipsoid, 
c. 0.5 — 0.7 mm x 0.5 mm; anthers exserted at male anthesis; infructescence 
stoutly oblong to tapering-cylindrical, 10 — 12 x 1.1 — 2.5 cm. 

Distribution: Papua New Guinea (Western Prov.), Solomon Islands 
(Guadalcanal, New Georgia, San Cristobal and the Santa Cruz Group), 
Fiji, Western and American Samoa. 

Habitat: Well-drained primary and secondary lowland to upper hill forest, 
on rocky, red soils. 30- 1000 m altitude. 

Notes: 1. Rhaphidophora spuria is the only entire-leaved Rhaphidophora 
indigenous to Fiji and Samoa. Examination of the types of R. spuria, R. 
graeffei and R. reineckei reveals no characters separating these species 
hitherto recognized as endemic for Fiji (R. spuria) and the Samoan islands 
(R. graeffei and R. reineckei). The earliest name, R. spuria, is adopted 
here. 

2. Rhaphidophora spuria is most similar to R. intonsa from Papua New 
Guinea (Morobe and Central Prov.) but may be readily distinguished by 
the truncate to weakly cordate lamina base and in having very much less 
fibre at the shoot tips. 

3. Despite the geographical disjunction, the single collection from mainland 
Papua New Guinea is of this species, matching the Pacific plants in 
vegetative and floral characters. 

Other specimens seen: PAPUA NEW GUINEA. Western Prov.: Fly River, Madiri Plantation, 
Womersley & Simmonds 5047 (GH). SOLOMON ISLANDS. New Georgia: Hovoro, 
Cowmeadow's collectors BSIP 3791 (BSIP, K, SING); San Cristobal: Ridge west of Warahito, 
Hunt RSS 2297 (BSIP, K, L); Santa Cruz Group: Vanikolo ( Vanikoro) Island, Kajewski 584 
(GH, K); Guadalcanal: adjacent to Tina River, 12 miles inland from coast, Womersley & 
Whitmore BSIP 1110 (BSIP, K, L, SING). FIJI. No further data, Copt. WUkes Exped. s.n. 
(US); Viti Levu: Nausori Highlands, 16 km from Bukika Sawmill, Melville & Melville 



Rhaphidophora in New Guinea. Australia and Western Pacific 



165 




Figure 30. Rhaphidophora spuria (Schott) Nicolson 

A. flowering shoot x 'A; B. leaf x 3 / 10 ; C. spadix detail, male anthesis x 10. A from Kajewski 584; 
B from Teraoka & Kennedy 64; C from Smith 8653. 



166 



Card. Bull. Singapore 53 (2001) 



71.998 (K); Naitasiri, Colo-I-Suva, Parham 11246 (K); MBA, vicinity of Nalotawa, eastern 
base of Mt Evans Range, Smith 4443 (US); Namosi, northern base of Korombasambasanga 
Range, in drainage of Wainavindrau Creek, Smith 8653 (K, L, P, US); Namosi, hills east of 
Wainikoroiluva River, near Namuamua. Smith 8947 (K. L, P, US). Kadavu (Kandavu), 
Namalata Isthmus region, Smith 29 (K, P, US); WESTERN SAMOA. Upolu: Lake Lanuto'o, 
Christophersen 402 (BISH, K); Above Vaipouli, Christophersen & Hume 1902 (BISH, K, P, 
US); Tapatapa, Cox 34 (K); Near Lotofaga. Lafulemu Ranch (property of Fay Ala'ilima). 
west of Fagatola River, c. 0.5 km NW of the bridge, Teraoka & Kennedy 64 (US); Near Mt 
Leou'e. Whistler 743 (B, US); Savai'i: Sa\ai\ua,Christophersen 2956 (BISH, K. P); La Vai. 
above Salailua Christophersen 3004 (BISH, US); Above Asau, Block 28, Whistler 1036 (B, 
US). AMERICAN SAMOA. Pago Pago, Meebold 8209 (K). 

27. Rhaphidophora stenophylla K. Krause 

Rhaphidophora stenophylla K. Krause, Bot. Jahrb. Syst. 49 (1912) 94. — 
Type: Papua New Guinea ('Kaiser Wilhelmsland'). East Sepik Prov., Djamu, 
22 April 1908, Schlechter 17581 (B, holot; P, iso). 

Figure 31 

Large, rather robust, semi-leptocaul, homeophyllous liane to unknown 
ultimate height; seedling stage very slender terrestrial to climbing, leaves 
arranged in two ranks; pre-adult plants clinging, slender, with very narrow 
leaves; adult shoot comprised of elongated, clinging, physiognomically 
unbranched, non-flowering stems and free lateral, leafy flowering stems; 
stems terete to strongly compressed, but not rectangular in cross-section, 
without cataphyll, prophyll and petiolar sheath fibre, internodes 1 — 3 x 
0.2 — 2 cm, separated by prominent almost straight, slightly corky leaf scars; 
flagellate foraging stem absent; clasping roots arising from nodes and 
internodes; feeding roots not observed; leaves spiro-distichous in mature 
plants; cataphylls and prophylls caducous; petiole deeply canaliculate, 2 — 
16 x 0.2 — 1 cm, apical geniculum small and very obscure, basal geniculum 
weakly defined; petiolar sheath very prominent, broadly winged and 
extending to apical geniculum, persistent almost through to leaf fall; lamina 
entire, linear-lanceolate to narrowly lanceolate or narrowly oblong- 
lanceolate, slightly falcate, 9 — 38 x 1.5 — 4 cm, subcoriaceous, base subacute 
to slightly decurrent, apex acute to briefly acuminate with a prominent 
tubule; midrib raised abaxially, sunken adaxially; primary venation pinnate, 
slightly raised abaxially, almost flush adaxially; interprimaries reticulate- 
parallel to primaries, almost indistinguishable from them, slightly raised 
abaxially, almost flush adaxially; secondary venation reticulate, hardly 
visible; inflorescence solitary, subtended by a ± fully developed foliage 
leaf; peduncle slightly laterally compressed to terete, 6 — 10 x c. 1 cm; spathe 
broadly canoe-shaped, stoutly long-beaked, 11 — 13 x 1.5 — 2.5 cm, stiffly 



Rhaphidophora in New Guinea. Australia and Western Pacific 



167 




Figure 31. Rhaphidophora stenophylla K. Krause 

A. adult shoot with flowering branch x l / 3 ; B. leaf lamina x V 2 ; C. venation detail x 3; D. leaf 
lamina x V 3 ; E. pre-adult clinging shoot x l / 4 ; F. pre-adult creeping shoot x z / 3 ; G. inflorescence, 
spathe fallen x 1: H. spadix detail, male anthesis x 6. A-C. E & F from Nicolson 1430; D. G-H 
from Streimann NGF 44247. 



168 



Card. Bull. Singapore 53 (2001) 



fleshy, yellow, caducous leaving a large scar at the base of the spadix; 
spadix cylindrical, slightly curved, long stipitate, 6 — 9 x 1.5 — 2.2 cm, inserted 
level to almost truncate on stipe, white; stipe slender to stoutly terete, 
2.5 — 3 x 3 — 0.4 cm; stylar region rhombohexagonal, 1 — 2 x 1 — 1.5 mm, 
conical; stigma punctiform, prominently raised, c. 0.3 mm diam.; anthers 
strongly exserted at male anthesis; infructescence not observed. 

Distribution: Papua New Guinea (including the Louisiade Archipelago). 

Habitat: Monsoon lower montane forest to submontane primary forest on 
moderate to steep slopes, sometimes on limestone. 20-940 m altitude. 

Notes: 1. Unmistakable by the long, narrow leaves, persistent winged 
petiolar sheath, long stipitate spadix and prominently raised stigma. There 
exist plants with broader laminas than typical (Brass 27691 & 28311 are of 
this broad-leaf type) that may be confused with Rhaphidophora discolor 
(also flowering on free shoots and with long-stipitate spadix), although the 
latter is readily distinguished by the broader leaf laminas, petiolar sheath 
degrading to strips and fibres, larger (13.5 — 19 x 2.2 — 4 cm) spadix and flat 
stigmas. 

2. Scindapsus schlechteri K. Krause is superficially similar in its narrow leaf 
lamina and persistent-winged petioles but differs, aside from a single ovule 
per ovary (and thus one-seeded fruits), by the sessile spadix and thicker 
leaf lamina with almost no visible venation. 

Other specimens seen: PAPUA NEW GUINEA. West Sepik Prov.: Telefomin, Sandaun. 
Hak Valley, Gentry transect on slope above Nenem, SE of Mianmin Airstrip, Frodin & 
Morren 3155 (K); East Sepik Prov.: vicinity of Malu & April river, Ledennann 8614 (B 
spirit); Central Prov.: Port Moresby, Koitaki (c. 30 km WNW of Port Moresby), Can 12218 
(BM, SING); Sogeri Plateau, 5-7 miles beyond Kokoda Trail Monument. 30 miles east of 
Port Moresby, Nicolson 1430 (K, L. P, SING, US); Milne Bay Prov.: Louisiade Archipelago. 
Rossel and Misima Island, Narian, Brass 27691 (GH, L); Abaleti, Brass 28311 (L); Morobe 
Prov.: Wau, Bulolo. Middle L.A., Streimann NGF 44247 (GH, K, LAE. SING. US). 

28. Rhaphidophora stolleana Engl. & K. Krause 

Rhaphidophora stolleana Engl. & K. Krause, Bot. Jahrb. Syst. 54 (1916) 
79. — Type: Papua New Guinea, East Sepik Prov., April River, 24 May 
1912, Ledermann 7382 (B, holo). 

Figure 32 

Moderate, somewhat robust, semi-leptocaul, homeophyllous (?) liane 



Rhaphidophora in Sew Guinea. Australia and Western Pacific 



169 




Figure 32. Rhaphidophora stolleana Engl. & K. Rrause 

A. adult shoot with flowering branch x l / 4 ; B. leaf lamina x V 3 ; C. venation detail x 3: D. 
inflorescence, spathe sectioned x 1V : : E. spadix detail, post-male anthesis x 8. All from 
Ledermann 7382. 



170 



Gard. Bull. Singapore 53 1 2(H) I) 



unknown ultimate height; seedling and pre-adult plants not observed; adult 
shoot architecture not completely known, but observed to comprise of greatly 
elongated, strongly pendent, physiognomically unbranched, leafy, non- 
flowering stems giving rise to abbreviated short, free, sympodial, flowering 
stems; stems smooth, climbing stems not observed, free stems weakly four 
angled to + terete in cross-section, occasionally with two close longitudinal 
keels along one side, dull pale yellow brown, without prophyll, cataphyll 
and petiolar sheath fibres, internodes to 13.5 x 1 cm, flowering shoots 
abbreviated, arising from just above the axils of leaves and twisting to 
present inflorescences upwards; flagellate foraging stems not observed; 
clasping roots arising singly from the nodes and internodes, slightly 
pubescent; feeding roots not observed; leaves very weakly spiralled, 
scattered; cataphylls and prophylls membranous, caducous; petiole shallowly 
canaliculate, 13 — 14 x 0.35 — 0.7 cm, smooth, with a well defined apical and 
large, slightly prominent basal geniculum; petiolar sheath very prominent, 
extending to apical geniculum, swiftly caducous leaving a slight scar; lamina 
entire, elliptic to oblong-elliptic, 20 — 29 x 8 — 13 cm, membranous, drying 
discolorous, adaxially mid-brown, abaxially pale orange-brown, base 
subovate, very briefly decurrent, apex acute to briefly acuminate, slightly 
falcate with a minute apiculate tubule; midrib slightly raised abaxially, 
sunken adaxially; primary venation pinnate, slightly raised abaxially; 
interprimaries parallel to primaries, occasionally slightly reticulated, slightly 
raised abaxially: secondary and tertiary venation reticulate, prominent 
abaxially in dried specimens; inflorescence solitary, subtended by a foliage 
leaf; peduncle terete, c. 6 x 0.3 cm; spathe elongate-cylindric, stoutly short- 
beaked, c. 10.2 x 1.7 cm, thinly stiff-coriaceous, duration unknown; spadix 
slender cylindrical, sessile, inserted very obliquely on to the peduncle, c. 8 
x 1.1 cm; stylar region rhombohexagonal, 1 — 1.2 x c. 1 mm, truncate; stigma 
slightly longitudinally elongate to punctiform, c. 0.4 x 0.25 mm; anthers 
exsertion not observed; infructescence not observed. 

Distribution: Papua New Guinea (East Sepik Prov.). Known only from the 
type. 

Habitat: Lowland riverine forest. 20-50 m altitude. 

Note: In its growth form, with long pendent stems with abbreviated flowering 
shoots arising in the leaf axils, Rhaphidophora stollcana resembles R. 
brevispathacea. The inflorescences are, however, quite different, with the 
spathe ovoid-cylindrical, 3 — 4 cm long, spadix slender cylindrical, 2.9 — 3.4 
cm long in R. brevispathacea v. the spathe c. 10.2 cm long, spadix c. 8 cm 



Rhaphidophora in New Guinea, Australia and Western Pacific 



171 



long in R. stolleana. To date, R. stolleana is known only from East Sepik 
and R. brevispathacea only from Mimika/Digul Province. 

29. Rhaphidophora versteegii Engl. & K. Krause 

Rhaphidophora versteegii Engl. & K. Krause. Nova Guinea 8 (1910) 248, 
Nova Guinea 8 (1912) 805: K. Krause & Alderw.. Nova Guinea 14 (1924) 
213. — Type: Indonesian Papua, Mimika/Digul border, near Sabang, 25 
June 1907. Versteeg 1308 (B. holo: BO, L, K. iso). 

Rhaphidophora ledermannii Engl. & K. Krause, Bot. Jahrb. Syst. 54 (1916) 
81; Hay. Aroids of Papua New Guinea (1990) PI. XIV, b, synon. nov. — 
Type: Papua New Guinea, West Sepik Prov. (Felsspitz), 7 Aug. 1913, 
Ledermann 12684 (B, lecto; selected here). Engler & Krause cited two 
syntypes, the other, Papua New Guinea, West Sepik (Felsspitz), Aug. 1913, 
Ledermann 12722, is missing from Berlin and presumed destroyed. 

Figures 33 & 34 

Robust, large, pachycaul. heterophyllous liane to 20 m; seedling stage not 
observed: pre-adidt plants shingling: adult shoot architecture comprised of 
clinging, physiognomically unbranched, densely leafy flowering stems; stems 
smooth, mid-green, with cataphylls and prophylls persistent and drying 
dark yellow, degrading into parchment-like remains, internodes 1—4 x 
0.4 — 2.5 cm, separated by very prominent slightly oblique leaf scars: 
flagellate foraging stem absent: clasping roots densely arising from the nodes 
and internodes. smooth and drying with parchment-like epidermis: feeding 
roots not observed; leaves distichous: cataphylls and prophylls chartaceous. 
degrading into strips of tissue and weak fibres at tips of flowering shoots; 
petiole deeply canaliculate. 8 — 48 x OA — 0.8 cm, smooth, with faint to rather 
prominent dark dense speckling, apical geniculum prominent, basal 
geniculum very large but not prominent; petiolar sheath extending to apical 
geniculum, broad, chartaceous, short-persistent, degrading to papery strips 
and sparse fibres, then falling; shingling lamina entire, cordiform, 2.5 — 7.5 
x 3—6.5, chartaceous, base cordate, posterior lobes overlapping, apex obtuse 
and minutely apiculate; adult lamina entire to slightly or extensively 
perforated, perforations elliptic to rhombic, extending from c. + to entire 
width of lamina on each side of the midrib, lamina ovate-elliptic to oblong- 
lanceolate or oblong-elliptic, slightly oblique, 33 — 57 x 9.5 — 27 cm, sub- 
membranous, drying pale grey-green to bright green, base rounded to acute, 
apex acute to acuminate; midrib prominently raised abaxially, ± flush 
adaxially: primary venation pinnate, prominently raised abaxially, weakly 



172 



Gard. Bull. Singapore 53 12001) 



so adaxially; interprimaries sub-parallel to primaries, less prominent than 
interprimaries, slightly raised abaxially and adaxially; secondary venation 
feebly reticulate to subtesselate; inflorescence very rarely solitary, usually 
several together, each subtended by a prominent chartaceous prophyll and 
one or more chartaceous cataphylls, the entire synflorescence emerging 
from a mass of dried, chartaceous cataphyll remains; peduncle slender to 
stout, terete, partially to completely obscured by cataphylls, 4 — 12 x 0.2 — 
0.8 cm; spathe slender canoe-shaped, hardly to stoutly beaked, 5 — 10.5 x 
1 — 2 cm, stiff-fleshy, very thick-walled (up to 1 cm at tip), yellow to yellow 
green, marcescent to early fruiting, eventually falling leaving a prominent 
scar; spadix stoutly cylindrical, sessile, inserted almost level on peduncle, 
3 — 9.5 x 1 — 1.5 cm, white at male anthesis; stylar region conical, mostly 
hexagonal in top view, 0.9 — 1.2 x 1 — 1.1 mm; stigma punctiform, very 
prominently raised, those at the tip of the spadix even more raised, 0.1 — 
0.2 x c. 0.3 mm, glossy, almost black in dried material; anthers exserted at 
male anthesis; infructescence stoutly oblong-cylindrical, 6—9 x 1.4 — 2 cm. 

Distribution: Indonesian Papua, Papua New Guinea (including New 
Ireland). Seemingly widespread but, based on known collections, perhaps 
not common. 

Habitat: Lowland to lower montane primary to secondary rain forest on 
clays and silts. Sea level to 1500 m altitude. 

Notes: 1. Rhaphidophora versteegii is most similar to R. spathacea in having 
clusters of inflorescences subtended and interspersed by chartaceous 
prophylls and cataphylls. It is readily identifiable, and distinguished from 
R. spathacea, by the distinctive pale grey-green colour of dried material 
and the form of the juvenile growth phase. Further, leaf lamina perforations 
occur in most (but not all) individuals of R. versteegii: laminas of R. 
spathacea are never perforated. The arrangement and shape of the juvenile 
shingling leaves of R. versteegii are unique in the genus although known 
from only one collection (Croat 5252). 

2. The type of R. ledermannii differs primarily in the perforation of the leaf 
lamina (R. ledermannii has profoundly perforated laminas) and as such 
appears to represent distinct species. However, the occurrence of perforated 
and unperforated laminas in different plants of the same species is not 
rare, e.g. R. puberula Engl., R. megasperma Engl, (see Boyce 1999, 2001) 
and its occurrence in R. ledermannii is not sufficient to warrant segregating 
the two species. Furthermore, a suite of characters - clustered inflorescences 
arising on clinging stems and subtended by chartaceous, later weakly fibrous 



Rhaphidophora in New Guinea, Australia and Western Pacific 



173 




Figure 33. Rhaphidophora versteegii Engl. & K. Krause 

A. flowering shoot x V 9 ; B. leaf lamina x V 3 ; C. venation detail x 3; D. juvenile shingling stage, 
adaxial view x 'A; E. juvenile shingling stage, abaxial viewx 7 2 ;F. inflorescence, spathe removed 
x 1 ; G. spadix detail, post-male anthesis x 8; H. spadix detail, early fruiting x 8; J. stylar region 
and stigma, side view x 4. All from Croat 52752. 



174 



Gard. Bull. Singapore 53 (2001) 



prophylls and cataphylls, stoutly cigar-shaped spathes, prominently raised 
and, compared with style diameter, large, glossy black stigmas and petioles 
with small speckles (tannin cells?) - leaves little doubt that one species is 
involved. 

3. Perforated leaf laminas occur in a number of otherwise unrelated 
Rhaphidophora species (e.g., R. foraminifera (Engl.) Engl., R. puberula, R. 
versteegii, R. pertusa (Roxb.) Schott) suggesting that while a useful diagnostic 
tool, lamina perforation cannot be used to circumscribe taxonomically 
meaningful groups within Rhaphidophora. 

Other specimens seen: INDONESIAN PAPUA. No locality, Docters van Leeuwen 11174 
(L); Mimika/Digul Prov. boundary: Lorentz River, near Bivakeiland. Pulle 44 (B spirit, L. 
K). Pulle 67 (L, K). PAPUA NEW GUINEA. East Sepik Prov.: vicinity of Wewak, along 
trail beyond Boys Town (reform school operated by S.V.D. Missionaries), Croat 52752 
(MO); Southern Highlands Prov.: Aria, near Veta, Aet 375 (L); Gulf Prov.: Baimuru. 
Vailala River, 70 km at 110 from Baimuru. 50 km at 318 from Kerema, Croft et al. LAE 
61251 (L, LAE); New Ireland: Lamet, NW Lavongai (New Hanover), 5km S of Lai Bay, 
Croft & Lelean LAE 65461 (E, GH, K, L, LAE, M). 

30. Rhaphidophora waria P.C. Boyce, sp. nov. 

Vicinitate in donatione inflorescentiae condita, Rhaphidophora waria R. 
spathaceae proxima est, sed foliorum laminis minoribus angustioribus magis 
rigidioribus venis lateralibus primatiis eis interprimariis clare inter se 
diversis, spadice stiptato, stigatibus profunde excavatis differt. — TYPUS: 
Papua New Guinea, East Sepik Prov., Ambunti, along Yapa (Hunstein) 
River, 1 Aug. 1966, Hoogland & Craven 10796 (L, holo; GH, K, L, LAE, 
CANB, iso). 

Figure 35 

Small but robust pachycaul, homeophyllous (?) liane to unknown ultimate 
height; seedling stage not observed; pre-adidt plants climbing with leaves 
arranged in two ranks and distinctive by the conspicuous yellow chartaceous 
petiolar sheaths; adult shoot architecture comprised of clinging, 
physiognomically unbranched, densely leafy flowering stems; stems with 
conspicuous smooth papery epidermis and cataphylls and prophylls 
persistent and drying dark yellow, then degrading into parchment-like 
remains and fibres, internodes 1 — 2.5 x 0.3 — 0.9 cm, separated by almost 
straight leaf scars; flagellate foraging stem absent; clasping roots densely 
arising from the nodes and internodes, minutely pubescent; feeding roots 
not observed; leaves distichous; cataphylls and prophylls thinly but stiffly 
chartaceous, eventually degrading into strips of tissue and fibres, particularly 



Rhaphidophora in New Guinea, Australia and Western Pacific 



1~5 




Figure 34. Rhaphidophora versteegii Engl. & K. krause 

A. flowering shoot x 7„: B. leaf lamina x C. venation detaiJ x 4: D. inflorescence, spathe 
removed x 2: E. spadix detail, pre-anthesis x 8. A from Versieeg 1308: B-E from Croft & Lelean 
LA E 65461. 



176 



Card. Bull. Singapore 53 12001) 



at tips of flowering shoots; petiole deeply canaliculate, 3 — 15 x 0.2 — 0.4 cm. 
smooth, apical geniculum weakly prominent, basal geniculum slightly less 
so and mostly obscured by cataphyll remains; petiolar sheath extending to 
apical geniculum, persistent, eventually degrading in strips leaving fibres; 
lamina entire, lanceolate to elliptic, falcate, 6 — 23 x 1.5 — 5 cm, stiffly 
coriaceous, apex acute, briefly acuminate; midrib raised abaxially, sunken 
adaxially; primary venation pinnate, raised abaxially. slightly less raised 
adaxially; interprimaries sub-parallel to primaries and less conspicuous; 
secondary venation tessellate-reticulate, raised abaxially, nearly flush 
abaxially; tertiary venation similar to secondary venation although more or 
less invisible; inflorescences several together, each subtended by a prominent 
chartaceous prophyll and one or more chartaceous cataphylls, the entire 
synflorescence emerging from a mass of cataphyll remains; peduncle stout, 
terete to laterally compressed, obscured by cataphylls, 2 — 4 x 0.3 — 0.35 
cm; spathe canoe-shaped, slightly stoutly beaked, 4 — 5.5 x 1.5 — 2 cm, stiff- 
fleshy, duration unknown; spadix cylindrical, stipitate, c. 3.2 x 0.9 cm, cream; 
stipe 2 — 3 mm; stylar region rounded-conical, almost circular in plan view, 
0.25 — 0.4 mm diam.; stigma deeply excavated, c. 0.2 mm diam.; anthers not 
exserted at male anthesis; infrnctescence not observed. 

Distribution: Indonesian Papua (Mimika Prov.) and Papua New Guinea 
(East Sepik Prov.). 

Habitat: Stunted forest and mixed heath forest on steep slopes. 200-580 m 
altitude. 

Notes: 1. Based on inflorescence presentation, Rhaphidophora waria is 
close to R. spathacea but differs in the smaller, narrower, much stiffer leaf 
laminas with the primary lateral and interprimary veins clearly differentiated 
from one another, the stipitate spadix and the deeply excavated stigmas. 

2. The specific epithet is taken from the Wasuk name 'waria' for the plant, 
which is recorded on the type specimen label. It does not refer to the 
Waria valley. 

Oilier specimen seen: INDONESIAN PAPUA. Mimika Prov.: Freeport Concession Area, 
Garcinia site on road, Johns 10418 (BO, K, MAN). 



Rhaphidophora in New Guinea, Australia and Western Pacific 



111 




Figure 35. Rhaphidophora waria P.C. Boyce 

A. flowering shoot x V 4 : B. leaf lamina x 2 / 3 ; C. venation detail x 3; D. juvenile creeping stage x 
V 3 ; E. inflorescence, spathe sectioned x 1: F. spadix detail, post-male anthesis x 8. All from 
Hoogland & Craven 10796. 



178 



Card. Bull. Singapore 53 (2001) 



Insufficiently known species 
Rhaphidophora dahlii Engl. 

Rhaphidophora dahlii Engl., Bot. Jahrb. Syst. 25 (1898) 8; K. Schum. & 
Lauterbach, Fl. Schutzgeb. Siidsee (1900) 211; Engl. & K. Krause in Engl., 
Pflanzenr. 37 (IV.23B) (1908) 33. — Type: Papua New Guinea, East New 
Britain, Gazelle Peninsula, Ralum, Jan. 1897, Dahl s.n. (Bf , holo). 

With no extant type, no spirit material in B (where the rich spirit collection 
is occasionally a source of type material in the absence of a dried type 
specimen), no specimens annotated by Engler or Krause (who presumably 
would have been familiar with, and have annotated, authentic material), 
and an ambiguous protologue, it is impossible to unequivocally ascribe the 
name R. dahlii to any known species. 

On the face of it, attempts to match Engler's protologue to known 
Rhaphidophora in the Bismarck Archipelago should be straightforward. 
There are only five Rhaphidophora species present on New Britain (R. 
conica, R. hayi, R. jubata, R. korthalsii and R. mima) and two (R. hayi and 
R. schlechteri) on nearby New Ireland. Of these, R. korthalsii and R. hayi 
may be immediately discounted since neither is remotely similar to the 
vegetative characters described for R. dahlii. Of the remaining four species, 
R. schlechteri can be ruled out because of its stipitate spadix (that of R. 
dahlii is described as sessile), and R. jubata because it is far too large 
(petiole to 85 cm long, lamina to 90 cm long, spathe 16 — 24 cm, spadix 
13 — 21 v. petiole to 14 cm, lamina to 24 cm, spathe c. 4 cm, spadix to 3 cm 
in R. dahlii). The remaining species, R. conica and R. mima, are both 
possible candidates, but there remain several problems, not least of which 
is that while in description rather similar to R. dahlii, R. conica and R. 
mima are clearly distinguishable from one another on characters such as 
spathe persistence and shape of the stylar region, characters that are either 
not or only scantily recorded by Engler for R. dahlii. Thus it is impossible 
to be certain if either or neither is attributable to R. dahlii. 

Excluded species 

Rhaphidophora amplissima Schott = Epipremnum amplissimum (Schott) 
Engl. 

Rhaphidophora carolinensis (Volk.) Fosberg = Epipremnum carolinense 

Volk. 

Rhaphidophora cunninghamii Schott = Epipremnum pinnatum (L.) Engl. 



Rhaphidophora in Sew Guinea. Australia and Western Pacific 



179 



Rhaphidophora koidzumii Kanehira = Epipremnum carolinense 
Rhaphidophora lovellae F.M. Bailey = Epipremnum pinnatum 
Rhaphidophora neocaledonica Guillaumin = Epipremnum pinnatum 
Rhaphidophora pinnata (L.) Schott = Epipremnum pinnatum 
Rhaphidophora pertusa var. vitiensis (Schott) Engl. = Epipremnum 
pinnatum 

Rhaphidophora vitiensis Schott = Epipremnum pinnatum 
Rhaphidophora zippcliana Schott = Amydrium zippelianum (Schott) 
Nicolson 

Acknowledgements 

Thanks are due to Linda Gurr. Pat Davis and Emmanuel Papadopoulos 
for skillfully executing the illustrations that accompany this article, and to 
Dr Alan Radcliffe-Smith for the Latin translations of the diagnoses. Thanks 
also to Dr Alistair Hay (NSW) who critically reviewed the manuscript and 
offered much useful advice and to Dr Dan Nicolson (US) for advice on 
nomenclature. 

References 

Boyce. P.C. 1998. The genus Epipremnum Schott (Araceae — 
Monsteroideae — Monstereae) in west and central Malesia. Blumea. 43: 
183—213. 

Boyce. P.C. 1999. The genus Rhaphidophora Hassk. (Araceae- 
Monsteroideae-Monstereae) in Peninsular Malaysia, and Singapore. 
Gardens' Bulletin Singapore. 51: 183 — 256. 

Boyce. P.C. 2000a. The genus Rhaphidophora Hassk. (Araceae- 
Monsteroideae-Monstereae) in the Southern and Western Indonesian 
Archipelago. Gardens' Bulletin Singapore. 52: 101 — 183. 

Boyce. P.C. 2000b. The genus Rhaphidophora Hassk. (Araceae- 
Monsteroideae-Monstereae) in the Philippines. Gardens' Bulletin 
Singapore. 53: 213 — 256. 

Boyce. P.C. 2001. The genus Rhaphidophora Hassk. (Araceae- 
Monsteroideae-Monstereae) in Borneo. Gardens' Bulletin Singapore. 53: 
19—74. 

Boyce. P.C. & J. Bogner. 2000. An account of neotenic species of 



180 



Gard. Bull. Singapore 53 (2001) 



Rhaphidophora Hassk. (Araceae-Monsteroideae-Monstereae) in New 
Guinea and Australia. Gardens' Bulletin Singapore. 52: 89—100 

Engler, A. & K. Krause 1908. Araceae-Monsteroideae. In A. Engler (ed.), 
Das Pflanzenreich 37 (IV.23B) 4 — 139. Engelman, Leipzig. 

Mayo, S.J., J. Bogner & P.C. Boyce. 1997. Genera of Araceae. Royal Botanic 
Gardens, Kew, U.K. 

Schimper, A.F.W. 1903. Guilds in Plant-geography upon a Physiological 
Basis. Pt 2, ch. 2: 192 — 206. (English translation by W.R. Fisher, revised 
and edited by P. Groom & LB. Balfour.) Clarendon Press, Oxford, U.K. 

Stearn, W.T. 1992. Botanical Latin, 4 th Ed. David & Charles, Newton Abbot 
& London, U.K. 

Strong, D.R. & Ray, T.S. 1975. Host tree location behavior of a tropical 
vine (Monstera gigantea) by skototropism. Science. 190: 804 — 806. 



australasica = 1 
brevispathacea = 2 
conica = 3 

cravenschoddeana = 4 
cryptantha = 5 
discolor - 6 
fortis = 7 
geniculata = 8 
gorokensis = 9 
guamensis = 10 



Index Exsiccatorum 

hayi = 11 
intonsa - 12 
intrusa = 13 
jubata - 14 
kokodensis = 15 
korthalsii = 16 
microspadix = 17 
mima = 18 
neoguineensis = 19 
okapensis = 20 



pachyphylla = 21 
pe trie ana = 22 
pilosa = 23 
schlechteri = 24 
spathacea = 25 
spuria = 26 
stenophylla - 27 
stolleana = 28 
versteegii - 29 
waria - 30 



Aet 375 = 29; Aet & Idjan s.n. = 6; Anderson 147 = 11; Ave 4330 = 16, 4736 
= 25 

Barker 145 = 16; Bateson 89, 91, 92, 93 = 18; Beccari PP 409 = 3, PP 443 = 
16, Blake 19756 = 1; Boden Kioss s.n. = 17; Boraule et al. BSIP 9141 = 24; 
Bourdy 244, 228 = 16; Bowden-Kerby 5983 = 25; Boyce 1447 = 5; Boyland 
399 = 1; Branderhorst 324 = 8; Brass 2794 = 24, 7100, 7380, 23847 = 25, 
27691, 28311 = 21, 28608 = 3, 28618 = 25, 28831 = 11 

Cabalion 2370 = 16; Carr 12218 = 21, 13237 = 15, 14313 = 12, 14863 = 20, 
s.n. = 15; Christophersen 402, 2956, 3004 = 26; Christophersen & Hume 



Rhaphidophora in New Guinea, Australia and Western Pacific 



181 



1902 = 26; Clemens 2066 = 4, 4637 = 12, 7779 = 12, 8064 = 4, 10662 = 8, 
11151, 41197 = 17; Coode 8037 = 25; Cowley s.n. = 1; Cowmeadow's 
collectors BSIP 3791 = 26; Cox 34 = 26; Craven & Schodde 14, 286, 375 = 4: 
Croat 52575, 52614 = 1, 52752 = 29, 52760 = 5, 52763 = 16, 52763A = 4. 
53764 = 25, 52787, 52788 = 8, 52792, 52793, 52800 = 16, 52803 = 8, 52808 = 
25. 52818 = 4, 52540 = 16, 52842 = 4, 52849 = 16, 52952 = 25; Croft & 
Lelean LAE 65461 = 29, NGF 34561 = 15; Croft et al. LAE 61251 = 29. 
LAE 61974 = 15; Curry 939 = 16 

Dissing 2776 = 16; Djibdja s.n. = 3; Docters van Leeuwen 10750 = 16, 
11048, 11174 = 29, Domstreich 77 = 4 

Essig & Martin LAE 55124 = 19; Evans 581 = 25 

Falanrum & Faimaw 6795 = 25; Flecker 9005 = 22; Forbes 323 - 25; Foreman 
& Katik LAE 59291 A = 18; Foreman & Vinas LAE 60099 = 25; Fosberg 
31986, 32572 = 25; Frodin & Morren 3155 = 27, 3209 = 6; Frodin et al. 2553 
= 24: Furtado s.n. = 3 

Gafui BSIP 17296 = 18; Gafui et al. BSIP 10215 = 18, BSIP 12575 = 24; 
Gjellerup 330 = 19; Graeffe 73 = 26, Gray 202, 1055, 2862 = 22; Green 
RSNH 1267 = 16; Green s.n. = 5; Guppy 196 = 18 

Hartley 13098 = 20; Henty & Streimann NGF 38835 = 19; Hind 246 = 22; 
Hollrung 372 = 19. 746 = 1; Hoogland & Craven 10796 = 30; Hoogland & 
Schodde 6943 = 21; Hosokawa 8334 = 16; Hunt RSS 2297 = 26, RSS 2398 = 
16. RSS 2722 = 18; Hyland 5608 = 1, 9162 = 22 

Isles & Vinas NGF 33829, NGF 34488 = 16 

Jacobs 9331 = 8; Johns 8197 = 16, 9967 = 25, 9796 = 16, 9970 = 8, 70593 = 
25, 10418 = 30, 70437 = 25 

Kairo 237 = 25. 757 = 17, NGF 44078 - 24; TCa/Vo c£ Streimann NGF 30715 
= 19; Kajewski 584 = 26, 530 = 16, 1891, 2162 = 18; Kalkman 3389 = 21; 
Kayima 154 = 25; Koidzumi s.n. = 25; 7^0/e c& 7377 = 25 

Lam 711 = 16; 763 = 19; 7074 - 24; Lauterbach 827 = 8; Lavarack & 
Lavarack NGF 31091 = 18; Gi//7/o» 'Triton 48' = 25; Leac/j NGF 34337 
= 6; Ledermann 6641 = 16, 7352 = 28, 7384 = 25, 5569 = 6, 5674 = 27, 9590 
= 6, 70247 = 24, 70379, 70526 = 16, 72654 = 29, 14336 = 25; Lelean & 
Streimann LAE 52541 = 12; Ludvigson 20 = 16 



182 



Card. Bull. Singapore 53 (2001) 



Mauriasi BSIP 8429, BSIP 12268, BSIP 13632, BSIP 13977 = 18; McKee 
1877 = 1; Meebold 8209 = 26; Melville 3693 = 1; Melville & Melville 71.998 
= 26; Meyer Drees 279 = 19; Millar NGF 9938 = 19; Moi et al. 196 = 8; 
Morrison s.n. - 16 

Nakisi & Babala BSIP 8222 = 24; Nicolson 926 = 3, 7355 = 16, 1386 = 25, 
1402 = 19, 1409 = 25, 1416 = 21, 1427 = 19, 1430 = 27, 1431 = 12, 1432 = 24, 
1473 = 12, 1475 = 6, 7477 = 24, 1480 = 19, 1483 = 16, 1496 = 14, 1498 = 16, 
7572 = 11, 7576 = 18, 7520 = 16, 7522 = 18, 7529, 7530 = 14, 1540 = 11, 1542 
= 16, 7552 - 3, 1564 = 12, 7565 = 19, 7567, 7573 = 16, 7577 = 11, 1580 = 25 

Parham 11246 = 26; Peekel 296 = 24; Pulle 44 = 29, 66 = 16, 67 = 29, 254 = 
2, 353 = 16; Pullen 949 = 16, 5996 = 24 

Raynal RSNH 16222 = 16; Reinecke 594 = 26; Ridsdale & Lavarack NGF 
30628, NGF 31600 = 14; Rijkers 14841 = 11; Rinehart 15568 = 10; Robbins 
870 = 9 

Sands 2779 = 16, 6276 = 13, 6846 = 14: Sands et al. 2044 = 24, 2384 = 11; 
Schlechter 7785 - 25, 76329 = 19, 16436 - 21, 77557 = 27, 78752 = 25, 78298 
= 19, 18308 = 24, 18678 = 17, 78888 = 24, 18985 = 1; Schodde 3935 = 24, 
4070 = 18; Seeman 655 = 26; Smith 29, 4443, 8653, 8947 = 26, 70789 = 1; 
Stone 2366 = 18, 4101 = 10; Storck 911 = 26; Streimann LAE 52789 - 19, 
LAE 52964 = 8, NGF 23963 = 20, NGF 28712 = 6, NGF 44247 = 27, NGF 
47654 = 25; Streimann & Kairo NGF 39190 = 7; Streimann & Womersley 
LAE 51847 = 16 

Takamatsu 40 - 16; Teona BSIP 6303 - 24; Teraoka & Kennedy 64 = 26 
Utteridge 79 = 8 

Vandenberg & Katik NGF 42372 = 24; Versteeg 1191 = 8, 7308 = 29; 
Versteegh BW 12647 = 23; Vinas 151 = 9 

Waterhouse 207, 778 = 18; Webb & Tracey 7066 = 11; Whistler 743, 1036 = 
26; White et al. NGF 1555 = 1; Whitmore 6204 = 16; Widjaja 6344 = 21; 
Capt. Wilkes Exped. s.n. - 26; Womersley & Simmonds 5047 = 26; Womersley 
& Sleumer NGF 13971 = 9, Womersley & Whitmore BSIP 1110 = 26; 
C.C.Y. Wong 278 = 16 



Zippelius s.n. = 25 



Rhaphidophora in New Guinea, Australia and Western Pacific 



183 



Index to species 

Names in bold are accepted, names in italics are synonyms, numbers refer 
to accepted name 



Cuscuaria spuria Schott 162 

Epipremnum multicephalum Elmer 130 

Pothos celatocaulis N.E. Br 128 

Monstera latevaginata 130 

Rhaphidophora apiculata K. Krause 158 

australasica F.M. Bailey 88 

apiculata 158 

brevispathacea Engl. & K. Krause 91 

buergersii Engl. & K. Krause 154 

celatocaulis (N.E. Br.) Alderw 128 

conferta K. Krause 153 

conica Engl 92 

copelandii Engl 130 

cravenschoddeana P.C. Boyce 96 

cryptantha P.C. Boyce & C. Allen 99 

dahlii 178 

discolor Engl. & K. Krause 103 

drepanophylla Alderw 154 

engleri Kaneh 158 

forbesii Rendle 158 

fortis P.C. Boyce 104 

geniculata Engl 108 

gorokensis P.C. Boyce 1 1 1 

graeffei Engl 163 

grandifolia K. Krause 130 

grandis Ridl 130 

guamensis P.C. Boyce 112 

hayi P.C. Boyce & Bogner 116 

hollrungii Engl 88 

iboensis K. Krause 89 

intonsa P.C. Boyce 119 

intrusa P.C. Boyce 120 

jubata P.C. Boyce 124 

kanehirae Hatusima 158 

kokodensis P.C. Boyce 127 

korthalsii Schott 128 

var. angustiloba Ridl. ex Engl. & 

K. Krause 128 



latifolia Alderw 130 

ledermannii Engl. & K. Krause 171 

maxima Engl 128 

microspadix K. Krause 136 

mima P.C. Boyce 138 

neoguineensis Engl 140 

novoguineensis Engl 140 

nutans Ridl 136 

obliquata Alderw 154 

okapensis P.C. Boyce & Bogner 144 

oreophila Engl. & K. Krause 103 

pachyphylla K. Krause 146 

palattensis Engl. & K. Krause 158 

palattensis Koidz 158 

palawanensis Merr 130 

pallidivenia Alderw 94 

peekelii Engl. & K. Krause 154 

peepla var. storckiana (Schott) Engl 163 

petrieana A. Hay 149 

pilosa P.C. Boyce 151 

reineckei Engl 163 

ridleyi Merr 130 

schlechteri K. Krause 153 

spathacea Schott 158 

spuria (Schott) Nicolson 162 

stenophylla K. Krause 166 

stolleana Engl. & K. Krause 168 

storckiana Schott 162 

tenuis Engl 128 

trinervia Elmer 130 

trukensis Hosokawa 130 

versteegii Engl. & K. Krause 171 

waria P C. Boyce 174 

wentii Engl. & K. Krause 108 



Gardens Bulletin Singapore 53 (2001) 185-189. 



Cytology of Two New Species of Musa (Musaceae) and 
Their Sectional Relationship 

K. JONG 1 AND G. ARGENT 



Royal Botanic Garden Edinburgh, Edinburgh. Scotland, UK. 
'Also, Dept. Plant & Soil Science, University of Aberdeen, Scotland, UK 

Abstract 

Cytological observations were made on two recently described new Bornean species of 
Musa L., M. monticola and M. suratii. Both have 2n = 20. Taxonomic and biogeographic 
implications are discussed. 

Introduction 

Two new Bornean species of Musa {Musaceae) were recently described by 
Argent (2000), M. suratii Argent and M. monticola [Hotta ex] Argent, 
both from Sabah. Malaysia. 

Their sectional placement, however, was unconfirmed. As cytological 
information has historically been of major value in helping to determine 
the sections and relationships in Musa taxonomy (Simmonds, 1962). 
chromosome preparations of these two species were made. 

Materials and Methods 

Seeds collected in the field were germinated in garden compost under 
glass at 28°C. Young seedlings were potted up for growing on. Root tips 
were pre-treated in alpha-bromonaphthalene for 22 h at 4.5°C or in 
paradichlorobenzene for 4 h at room temperature (c. 20°C), fixed in 3:1 
ethanohglacial acetic acid and stored in this fixative until required. After 
hydrolysis in 5M HC1 at room temperature for 30 min, the roots were 
transferred to the Feulgen Reagent (prepared according to Fox, 1969) for 
2 h. After washing with tap water, roots were placed in an 1:1 enzyme 
mixture of 2% pectinase and 2% cellulase for 60 min. Squashing was done 
in 45% acetic acid or in 0.4% aceto-carmine. Permanent slides were 
prepared according to a modified freezing method outlined in Jong (1997). 



186 



Card. Bull. Singapore 53 (2001) 



Cytological Observations and Discussion 

Musa has a range of basic chromosome numbers that is associated with the 
traditional sections (Stover and Simmonds 1987) as follows: 



Section Basic Number 

Musa (formerly Eumusa) Baker x = 11 

Rhodochlamys Sagot x = 11 

Australimusa Cheesman x = 10 

Callimusa Cheesman x = 10 

[Musa beccarii] Incertae Sedis Simmonds x = 9 

Ingentimusa Argent x = 7 



One section, Ingentimusa is represented only by a single species, M. 
ingens Simmonds. The genus Ensete Horan has a basic number of 9, the 
same as Musa beccarii N.W. Simmonds, although there is no question that 
that species belongs with Ensete, its position within Musa is still problematic 
since molecular and morphological data (Jarret & Gawel 1995) appear to 
be in conflict. In fact, there has been a progressive tendency to amalgamate 
sections Musa and Rhodochlamys (Simmonds 1962, Jarret and Gawel 1995, 
Shepherd 1999). Further, Jarret and Gawel (1995) imply that Musa coccinea 
Andr. is sufficiently different from the other Musa species to be placed in 
a section of its own. The only basic number missing from this dysploid 
series of x = 7 to 11 is x = 8. 

The somatic chromosome number of both M. suratii and M. monticola 
is 2n = 20 (Fig. la-Id). There are, however, certain cytological differences 
between the two species that are worth noting. The chromosomes of M. 
suratii are larger than those of M. monticola, and at least two pairs of 
chromosomes bear satellites. There is also a gradual gradation of 
chromosome size not obvious in M. monticola, where only one satellited 
chromosome pair is visible. Interphase nuclei are finely granulose, with no 
distinctively stained chromocentres, in contrast to those of M. monticola, 
where chromocentres are obvious and numerous. Unlike M suratii, there 
is a characteristic tendency for chromosomes in M. monticola to associate 
in twos, often so closely together that they might be mistaken as single 
chromosomes; quite frequently there is one association of three as illustrated 
in Fig lc and Id. 

In terms of chromosome number, both new species fall within either 
sect. Callimusa or sect. Australimusa whose basic chromosome number is x 
= 10. At present we have insufficient evidence to say whether the difference 
in the chromosome morphology and behaviour noted above are consistent 



Cytology of two new Musa Species 



187 






\ ' 


* 


*u * 


b 





• 




Figures la and lb Musa suratii. metaphase. 2n = 20. lb is a drawing of la, solid arrowheads 
indicating satellited chromosomes. Open arrow points to chromosome partially out-of-focus. 
Figures lc and Id Musa monticola, metaphase. 2n = 20 from different roots. Note marked 
close association of certain chromosomes in twos, and one in a group of three. Open arrow 
points to out-of-focus chromosome. Scale bar = 10 urn 



or taxonomically significant between these two sections. 

Section Callimusa has distinctive seed morphology with elongated, 
barrel-shaped seeds that have a large oil store, which is represented in old 
seed by an air space. Most of the species also have well-developed 
corrugated, scarious auricles at the apex of the leaf sheath ("shoulder' as 
defined by Argent 1976). Australimusa seeds by contrast are variable from 
globose to highly angled, the leaves have variable 'shoulders' on the leaf 



188 



Card. Bull. Singapore 53 (2001) 



sheaths, which may occasionally be loosely corrugated but are never 
expanded into auricles. On these morphological grounds, both species are 
clearly associated with sect. Australimusa rather than sect. Callimusa. 

Hotta (1987) clearly realised the relationship between Musa monticola 
and the local form of M. textilis Nee in Sabah and the seed, although 
slightly larger and more distinctly mammilose, is very similar to that of 
cultivars of M. textilis. Musa suratii has the smallest seed recorded in the 
genus but the morphology is even closer to that of M. textilis, which is the 
type species of sect. Australimusa. However, Musa suratii has extraordinary 
morphology with green flowers, orange bracts, a very slender habit and 
copious wax. All of these characters considerably expand the concept of 
sect. Australimusa. The description of two more species in sect. 
Australimusa from Borneo moves the emphasis of the distribution a little 
more to the west as previously five of the six known species occurred in 
New Guinea and the Solomon Islands. Now with the uncertain status of 
M. muluensis Hotta and M. tuberculata Hotta, it is looking as if Borneo 
may actually encompass more variation in this section than is found in 
New Guinea. 

Acknowledgements 

We wish to thank all the horticultural staff who have looked after the 
living banana collections, in particular Mr D. Mitchell and Mr P. Smith 
who have them in their overall charge. Mary Mendum has provided 
important technical assistance for which we are most grateful. 

References 

Argent, G.C.G. 1976. The wild bananas of Papua New Guinea. Notes 
from the Royal Botanic Garden Edinburgh. 35: 77-114. 

Argent, G. 2000. Two interesting wild Musa species (Musaceae) from 
Sabah, Malaysia. Gardens' Bulletin Singapore. 52: 203-210. 

Fox, D. 1969. Some characteristics of the cold hydrolysis technique for 
staining plant tissues by the Feulgen reaction. Journal Histochemistry 
and Cytochemistry. 17: 226-232. 

Hotta, M. 1987. Distribution of the genus Musa in Malaysia. Acta 
Phytotaxonomica et Geobotanica. 38: 292-302. 

Jarret, R.L. and N. Gawel. 1995. Molecular markers, genetic diversity and 



Cytology of two new Musa Species 



189 



systematics in Musa. In: Gowen, S. (ed.). Bananas and Plantains. 
Chapman & Hall, U.K. Chapter 3. 

Jong. K. 1997. Laboratory Manual of Plant Cytological Techniques. Royal 
Botanic Garden Edinburgh, Edinburgh, U.K. vi+96 pp. 

Shepherd, K. 1999. Cytogenetics of the Genus Musa. International Network 
for the Improvement of Banana and Plantain, Montpellier, France. 

Simmonds, N.W. 1962. The Evolution of the Bananas. Longmans, U.K.. 

Stover. R.H. and N.W. Simmonds.1987. Bananas. 3 rd ed. Longmans Scientific 
& Technical, U.K. 



Gardens' Bulletin Singapore 53 (2001) 191-239. 



The Genus Etlingera (Zingiberaceae) in Peninsular 
Malaysia Including a New Species 

S.H. KHAW 

23 Jalan SS2/81 
47300 Petaling Java. Selangor. Malaysia 

Abstract 

Etlingera pieeae Khaw from the Temenggor Forest Reserve in Perak. Peninsular Malaysia, 
is described as a new species. Including this new species, there are 12 Etlingera species in 
Peninsular Malaysia for which a key and descriptions are provided. 

Introduction 

Etlingera species are recognized by the presence of an involucre of large, 
sterile bracts, elongate and tubular bracteoles (Fig. 1). flowers with a distinct 
tube formed by the bases of the labellum and the filament above the 
attachment of the corolla lobes (Fig. 2). and the labellum inrolling as the 
flower withers (Burtt and Smith, 1986). 

The first comprehensive systematic account of Peninsular Malaysian 
species now included in Etlingera was given in the classic monograph of 
Holttum (1950). His detailed account included four Phaeomeria species and 
six Achasma species, now all considered as species of Etlingera (Table 1). 

Holttum (1950) credited Valeton for recognizing that Phaeomeria is 
closely related to Achasma. The two major characters that Holttum (1950) 
used to distinguish between these two genera were the peduncle (long and 
erect in Phaeomeria, short and usually subterranean in Achasma) and the 
labellum (short in Phaeomeria and elongate in Achasma). 

Subsequently, Burtt and Smith (1986) united the two genera Achasma 
and Nicolaia Horan., which replaced Phaeomeria, an invalid name (Burtt 
and Smith, 1986) with a third, Geanthus Valeton, to form a composite 
genus, Etlingera Giseke. (Geanthus is not found in Peninsular Malaysia). 
In uniting the three into a single composite genus, Burtt and Smith (1986) 
maimained that the two characters conventionally used to divide Achasma, 
Geanthus and Phaeomeria into distinct genera were inadequate. In addition, 
they observed that the anther is held at an angle to the filament in Achasma. 
whereas it is erect in Geanthus and Phaeomeria. 



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Card Bull. Singapore 53 (2001) 



Table 1. Etlingera species in Peninsular Malaysia, including the names 
used by Holttum. 



Etlingera species 


Name in Holttum (1950) 


t. elatior 


Phaeomeria speciosa 


E. maingayi 


P. maingayi 


E. julgens 


r. julgens 


E. venusta 


/ . venusta 


E. pauciflora 


Achasma pauciflorum 


E. punicea 


A. macrocheilos 


E. subtevraneci 


A. subterraneum 


E. metriocheilos 


A. sphaerocephalum 


E. triorgyalis 


A. trior gy ale 


E. littoralis 


A. megalocheilos 


E. comer i 




E. pieeae 





Further to Holttum's enumeration of the species, while making an 
inventory of gingers of the Temenggor Forest Reserve, Perak, Peninsular 
Malaysia, in 1994 and 1995, two unnamed species of Etlingera were collected 
(Khaw, 2000). One, E. corneri J. Mood & H. Ibrahim, was recently described 
(Mood & Ibrahim 2000); the other, E. pieeae Khaw, is described here. 

The new species, E. pieeae, is particularly interesting as it is in some 
respects intermediate between Holttum's Phaeomeria and Achasma. In 
Peninsular Malaysia, the two are easily recognized based on inflorescence 
characters (the flower head of the Phaeomeria group being either cup- 
shaped on a long, aerial stalk or in the Achasma group sub-cylindric on a 
short, subterranean stalk). However, the inflorescences of E. pieeae do not 
have the typical appearance of either group. Instead, because its sterile 
bracts are not broad but oblanceolate, spreading slightly and loosely 
imbricating at the bases, a wide cup-like or a narrow sub-cylinder of sterile 
bracts is not obvious. In this aspect, its inflorescence resembles that of E. 
pauciflora (formerly a species of Achasma), which has the typical 
subterranean inflorescence. In addition, E. pieeae is intermediate between 
the two groups in the flower head having a relatively long peduncle. The 
result is that the distal part of its peduncle together with the flower head 
are aerial, a character of the Phaeomeria group. These features, coupled 
with flowers with a long labellum, (a character of Achasma) make it distinct 
from other Eltingera species in Peninsular Malaysia. The fact that it is 
intermediate between the two groups adds support for uniting Phaeomeria 



Etlingera I Zingiberaceae ) in Peninsular Malaysia 



193 



and Achasma into a single genus. 

Illustrated accounts of Peninsular Malaysian Etlingera were given by 
Weber (1995), Larsen et al (1999) and Lim (2000). 

The attractive and colourful inflorescences and infructescences of 
Etlingera species are of horticultural potential. In Peninsular Malaysia, 
only kantan, E. elatior (Jack) R.M. Smith, is grown as an ornamental and 
commercially for culinary purposes. The rest are not widely cultivated 
horticulturally or for the cut-flower trade, although they are gaining 
popularity in subtropical Australia and U.S.A.. 

Materials and Methods 

This account is based on the examination of the living specimens. Field 
observations on the morphology of the living plants and the development 
of the inflorescence and infructescence were made for all the Etlingera 
species, except for E. subterranea and the fruits of E. littoralis, for which 
material was not available. In addition, observations were made of plants 
cultivated from rhizomes and seed collected from the field. Material fixed 
in Copenhagen mixture, colour photographs, as well as dried specimens 
were prepared for study. Observations of fresh materials, unless otherwise 
stated, form the basis of the characters used in the key and the descriptions 
of the species. Drawings of dissected Etlingera flowers follow the style of 
Burtt and Smith (1986). Herbarium material from the Etlingera collections 
at KEP. KLU and SING were examined. Types not seen are indicated 
(5.V.). Photographs of the type specimen of E. venusta at K were made by 
Dr Saw Leng Guan. 

Compared with Holttum's account, several additional characters have 
proved useful in identifying the species, such as the production of stilt 
roots, number of flowers, the aspect of the flower head in lateral view, 
shape of the basal lobes of the labellum. the form of the epigynous glands 
and the scent of the crushed leaf sheaths. Examination of the minute 
epigynous glands that Holttum (1950) called stylodes (nectary glands) 
reveals that there are two main types in Etlingera (Fig. 3). the one with a 
rounded or irregular apex is found in the Phaeomeria group of species, the 
other type with a narrowed apex with the sole exception of E. metriocheilos 
is found in the Achasma group. 

The description of the fruit of E. metriocheilos in the present work 
differs from that of Holttum. He described the fruit as smooth based on 
the examination of a single specimen {Ridley s.n. Maxwell's Hill. June 
1893). whereas examination of living material shows the fruits to be ridged 
longitudinally, each ridge with a shoulder-like apical mound, covered with 




Figure 1. Inflorescence types in Etlingera. A, E. maingayi; B. E. elation C, E. fulgens; D. E. 
triorgyalis; E, E. pauciflora; F. E. punicea; G, E. metriocheilos; H, E. littoralis; (from living 
material). 




Figure 2. Flower and labellum types in Etlingera. A, B, E. maingayi; C, D, E. elation E, F, E. 
fulgens; G, H, E. triorgyalis; I, J, E. pauciflora: K. L, E. methocheilos: M, N, E. punicea: O. P, E. 
littoralis; (left, flower in lateral view; right, dissected flower; from spirit material). 



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Figure 3. Epigynous glands in Etlingera. A, E. maingayi; B, E. fulgens; C, E. elation D. E. corneri; 
E. E. venusta; F, E. pieeae; G, E. triorgyalis; H. E. pauciflora: I, E. punicea; J, E. metriocheilos; K. 
E. littoralis: (A-K. left, adaxial view; right, abaxial; from spirit material). 



Etlingera (Zingiberaceae) in Peninsular Malaysia 



199 




0.2 cm 



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Card. Bull. Singapore 53 (2001) 




Figure 4. Infructescence types in Etlingera. A, E. maingayi; B, E. elatior; C, E. fulgens; D, E. 
triorgyalis; E, F, G, E. pauciflora; H, E. punicea; I, £. metriocheilos; (from spirit material). 



Etlingera IZingiberaceae) in Peninsular Malaysia 



201 




202 



Gard. Bull. Singapore 5.? (2001 ) 



prominent warts (Fig. 41). In addition, infructescences of three species (E. 
trior gyalis, E. pauci flora and E. punicea) stated as unknown by Holt turn, 
are described and illustrated here (Fig. 4D, E-G, H). 

Description of the Genus Etlingera in Peninsular Malaysia 

Rhizomes at, just below, or deeper in the ground (E. pauciflora, E. punicea 
and E. littoralis); elements long and slender between the leafy shoots in 
the smaller species (E. maingayi and E. pauciflora) or stout in the larger 
species; with stilt-roots above the soil, leaf-litter or swampy ground (E. 
maingayi and E. corneri). Leafy shoots tall, to 6 m in the larger species, 
often with swollen, bulbous bases; close together forming clumps or widely 
spaced to about 1 m apart; sheaths ribbed longitudinally with cross-bars 
covered by tufts of short, white hairs (often clearer on young sheaths); 
scent of crushed sheaths either a pleasant sour scent (all species in the 
Phaeomeria group and including E. corneri) or strong and peppery (in E. 
triorgyalis and E. pieeae) or without a distinct odour; ligule bilobed or 
entire; petiole usually present or lamina subsessile (E. venusta); laminas 
many, large, to about 1 m long, often when young pink or with brownish- 
pink broad bars or deep purple-red beneath (E. fulgens and E. 
metriocheilos). Inflorescence on a separate leafless shoot, arising from the 
base of the leafy shoot. Peduncle covered by bladeless sheaths in two 
ranks, widely spaced and not overlapping (the Phaeomeria group) or 
overlapping (the Achasma group); long, slender with a short horizontal 
subterranean axis, which turns upwards, forming a much longer, erect, 
aerial stalk (Fig. 5A) with the flower head raised well above the ground (in 
the Phaeomeria group and E. corneri) or with the horizontal subterranean 
part of the peduncle relatively longer (Fig. 4D), the erect part only a few 
cm long with the flower head just below ground level, so that only the 
opened flowers are displayed on the soil surface (the Achasma group) or 
the peduncle is at first horizontal and subterranean but then erect and 
producing a flower head well above the ground level (E. pieeae. Fig. 7 A, 
B). The flower head enclosed by broad sterile bracts; usually tightly 
imbricating and forming either a capitate, wide cup-like structure (the 
Phaeomeria group. Fig. 1 A-C, 5B, 6B) or a narrower, sub-cylindric structure 
(the Achasma group. Fig. ID, F-H). In both cases, in side view the sterile 
bracts hide a portion of the flower-head proper except in E. pauciflora, E. 
subterranea and E. pieeae where the narrow lanceolate sterile bracts are 
loosely imbricating at the bases only, so do not form a sub-cylindric structure 
(Fig. IE, 7B), their inflorescence heads proper thus in side view are visible 
between the bracts. Receptacle of inflorescence with apex inflated, rounded 



Etlingera I Zingiberaceae ) in Peninsular Malaysia 



203 



or flat, usually elongating only a few cm but up to 17 cm in E. elatior; with 
numerous (55 — 200 or more) flowers per flower head in the Phaeomeria 
group, 30 — 60 in the Achasma group, 20 — 35 in E. pieeae and 1 — 3 flowers 
in E. pauciflora and E. subterranea. Sterile and fertile bracts: the outermost 
sterile bracts show a transition from the uppermost sheath of the peduncle, 
bracts usually not persisting until fruit maturity, coloured or whitish, erect 
or spreading, much broader than the fertile bracts: outermost fertile bracts 
show a transition from the sterile bracts, decreasing in size towards the 
centre of the flower-head: each fertile bract producing one flower. Bracteole 
tubular, two-lobed. deeply slit down the opposite side. Calyx tubular. 3- 
lobed. deeply slit down the opposite side. Corolla tubular, often densely 
hairy at entrance to tube; with three erect lobes (petals), the dorsal lobe 
slightly wider than the lateral ones. Labellwn with a short or elongated 
blade, broadening at the base to form distinct basal lobes (Fig. 2, 7H) or 
without basal lobes (E. venusta and E. corneri. Fig. 5H. 6J): the lowest part 
of the labellum joined to the base of the stamen to form a short, fleshy 
staminal tube above the bases of the corolla-lobes: usually with the basal 
lobes erect on either side of the anther or folded together, partially or 
entirely covering the anther; after flowering, the lower part of the blade 
inrolling while the distal part withers; with the blade erect or spreading 
horizontally and the apex slightly reflexed; the apex entire or bilobed. 
Staminodes lacking. Filament short or in E. venusta and E. corneri almost 
sessile. Anther erect or bent forward towards the labellum. Epigynous 
glands as two fleshy, rounded mounds surrounding the base of the style, 
each mound irregularly lobed at the apex in the Phaeomeria group (Fig. 
3A-E) and E. metriocheilos (Fig. 3J) or as two fleshy, compressed blades, 
not encircling the base of the style, each blade not further lobed. apex 
shortly pointed in most species of the Achasma group (Fig. 3F-I, K). Ovary 
glabrous or hairy, stigma rather large. Infructescence large, usually globular 
but elongate in E. elation fruits close together, pericarp smooth and each 
with a long persistant calyx in the Phaeomeria group (Fig. 4A-C. 5F 6M) 
or ridged longitudinally and with warts in the Achasma group (Fig. 4D. H, 
I) or in E. pauciflora (Fig. 4E-G) and E. littoralis (Holttum. 1950) smooth; 
indehiscent. fleshy or dry: maturing within about two months. Seeds 
irregularly angular and truncate, each surrounded by thin, whitish, 
translucent pulp. 



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Card. Bull. Singapore 53 (2001) 



Key to Etlingera species in Peninsular Malaysia 

la. Inflorescence raised well above the ground on peduncles 40-200 cm; 
sterile bracts showy, aerial, tightly imbricating, with their upper half 
spreading horizontally, forming a very shallow or deep, wide, cup 
(with diameter exceeding or nearly the same as the height); lip with 
a very short blade 2 

lb. Inflorescence almost entirely subterranean on peduncles 5-20 cm or 
rarely (only in E. pieeae) raised above the ground on peduncles to 
48 cm long; sterile bracts not showy, usually embedded in the ground 
except for their distal parts, usually tightly imbricating, then forming 
a long narrow sub-cylinder (with diameter always less than the height) 
or loosely imbricating at the bases only, thus not forming a sub- 
cylinder; lip with elongated blade 6 

2a. Sterile bracts tightly imbricating at the bases only, forming a very 
shallow, inconspicuous cup, with the upper half of the bracts 
spreading horizontally; receptacle of inflorescence elongating 
markedly during flowering forming a cone-shaped flower head within 
the involucral cup 1. E. elatior 

2b. Sterile bracts tightly imbricating, forming a firm cup; receptacle of 
inflorescence hardly elongating during flowering forming a flat or 
bowl-like flower head at side view 3 



3a Involucral cup up to c. 3 cm high and c. 4 cm wide, in side view the 
cup hiding a third of the flower-head; sterile bracts densely appressed 
white-hairy all over surface 2. E. maingayi 

3b. Involucral cup up to c. 9 cm high and c. 7.5 cm wide, in side view the 
cup hiding almost entirely the flower-head; sterile bracts glabrous or 
only sparsely white-hairy 4 

4a. Leaves with upper surfaces glossy, dark green, lower surfaces 
conspicuously purple-red when young; lamina smooth, not 
prominently ribbed, margin markedly crisped; sterile bracts 
subrotund, strongly incurved at apex; lip with blade widening at 
base to form distinct basal lobes, red with yellow margin; fruits pale 
green, densely appressed short-hairy 3. E. fulgens 

4b. Leaves entirely green on both surfaces even when young; lamina 
prominently ribbed, margin not markedly crisped; sterile bracts 
oblong-elliptic, recurved at apex; lip with blade narrowed to base, 
not widening to form distinct basal lobes, white with red median 
band; fruits bright red and glabrous 5 



Ellingera (Zingiberaceae) in Peninsular Malaysia 



205 



5a. Stilt-roots absent; petiole always inconspicuous (leaf subsessile); leaf 
base always cuneate: sterile and fertile bracts with acute apices, 
purple-pink, pale green at the base; the involucral cup completely 
decayed before fruit maturity 4. E. venusta 

5b. Stilt-roots 10-25 cm long; petiole 1.5^1 cm long; leaf base mostly 
rounded (sometimes cuneate); sterile and fertile bracts with 
subtruncate apices, bright red. white at the base; involucral cup 
persisting to fruit maturity 5. E. corneri 

6a. Peduncles to 30 cm; fertile part of inflorescence with all sterile bracts 
raised well above the ground; bracts loosely imbricating at the bases 
only, not forming a sub-cylinder, in side view the flower head visible 
between the bracts 6. E. pieeae 

6b. Peduncles 3-15 cm; fertile part of inflorescence always subterranean 
except for distal parts of sterile bracts; bracts tightly imbricating, 
forming a long, narrow sub-cylinder, in side view hiding one to two 
thirds of the flower-head 7 



7a. Flowers 1-3 per inflorescence 8 

7b. Flowers 30-60 per inflorescence 9 

8a. Corolla lobes 1-2 cm longer than calyx, dorsal lobe 3-4 x c. 1.5 cm; 

lip yellow at centre with red margins 7. E. pauciflora 

8b. Corolla lobes almost same length as calyx, dorsal lobe 1-1.5 x c. 0.5 

cm; lip crimson with white margins 8. E. subterranea 

9a. Corolla c. 2 cm longer than calyx; dorsal corolla lobe hiding the 
entire anther; stigma hidden under dorsal corolla lobe; lip with yellow 
median band and red margins 9. E. punicea 

9b. Corolla about the same length as calyx; dorsal corolla lobe hiding 
only the basal part of anther; stigma clearly visible; lip entirely red 
or with yellow, orangey-red or white margins, always without a yellow 
median band 10 



10a. Leaf sheaths purple-red; lower surface of young leaves purple-red; 
sterile bracts deep-red; lip purple-red with white margin; stigma 
purple-red, almost black 10. E. metriocheilos 

10b. Leaf sheaths green; lower surface of young leaves entirely green or 
green flushed pale-pink; sterile bracts white or pale green with pink 
apices; lip entirely red or with a yellow margin; stigma pale-pink or 
bright red 11 



206 



Card. Bull. Singapore 53 (2001) 



11a. Outer sterile bracts with apices recurved; lip entirely pink-red; stigma 
pale-pink; crushed leaf sheaths with a distinct peppery scent; inner 
surface of staminal tube densely hairy and conspicuously warty; fruits 
with longitudinally ridged pericarp 11. E. triorgyalis 

lib. Outer sterile bracts with apices not recurved; lip entirely red or with 
a yellow or orange-red margin; stigma bright red; crushed leaf sheaths 
without any distinct scent; only inner surface of staminal tube densely 
hairy, not warty; fruits with smooth pericarp 12. E. littoralis 

Description of the Species 

/. Etlingera elatior (Jack) R.M. Sm., Notes Roy. Bot. Gard. Edinb. 43 
(1986) 244; Lim, Folia malaysiana 1 (2000) 4. Syn.: Alpinia elatior Jack, 
Misc. mal. 3, 7 (1822) 2 (s. v.); Elettaria speciosa Blume, Enum. PI. Javae 
(1827) 51; Alpinia magnified Roscoe, Monandr. PI. (1828) t. 75 (s.v.); 
Phaeomeria imperialis Lindl., Nat. Syst. Ed. 2 (1836) 446 (s.v.); Ridl., Fl. 
Malay Penins. 4 (1924) 272; Nicolaia imperialis Horan.. Prodr. Monogr. 
Scitam. (1862) 32, t. 1. (s.v.); Nicolaia speciosa Horan., Monogr. Scitam. 
(1862) 32 (s.v.); Valeton, Bull. Jard. Bot. Buitenz. 3 rd Ser. 3 (1921) 138; 
Hornstedtia imperialis Ridl., in J. Str. Br. Roy. Asiat. Soc. 32 (1899) 148; 
Mat. Fl. Malay. Penins. (1907) 40; Phaeomeria magnified (Roscoe) K. 
Schum., Pflanzenr. Zingib. (1904) 262; Phaeomeria speciosa (Blume) Merr., 
Enum. Philip. PI. 1 (1922) 241; Holttum, Gard. Bull. Sing. 13 (1950) 181; 
Henderson, Malayan Wild Flowers, Monocots. (1954) 153. Type: Sumatra, 
Jack s.n., (s.v.) - specimen lost? 

Figures IB, 2C & D, 3C, 4B 

Rhizomes just below ground; stout, 3^4 cm diam.; scale-leaves overlapping, 
green. Leafy shoots 5-6 m tall; 10-18 cm apart, forming clumps; young 
sheaths longitudinally ribbed, with cross-bars not covered by tufts of short, 
white hairs; crushed sheaths with a pleasant, sour fragrance; ligitle c. 2 cm 
long, apex broad, entire or slightly bilobed, margin with short brown hairs; 
petiole 2.5-3.5 cm long, surface longitudinally ribbed. Laminas c. 17 pairs; 
(topmost) c. 33 x 4.5 cm, (middle) 66-81 x 15-18 cm, (basal) c. 36 x 14 cm; 
upper surface with slightly raised lateral veins; entirely green on both 
surfaces, (sometimes flushed pink in young leaves); lower surface glabrous; 
base variable, mostly broadly rounded or cordate, sometimes cuneate or 
unequal. Peduncle to 100-200 cm; shortly horizontal, then erect, aerial, 
close to base of leafy shoot; sheaths c. 13, shortest at base 5-20 cm long, 
apex rounded with subapical tooth 0.1 cm long, pale green at apex, dark 
green towards base; not overlapping, widely spaced. Inflorescence raised 



Etlingera ( Zingiberaceae) in Peninsular Malaysia 



207 



well above the ground. Fertile apical part of inflorescence with well 
developed sterile bracts tightly imbricating at the bases only, forming a 
very shallow inconspicuous cup-like structure c. 2.5 cm high. 6 cm wide 
and with the upper half of the bracts reflexed: in side view the cup hiding 
c. 1/3 or less of the conical inflorescence head proper: the cup not persistent 
until fruit maturity. Receptacle of inflorescence 4-9 cm long, lengthening to 
17 cm (usually shorter): apex inflated and rounded: bearing up to 200 or 
more flowers: 11-13 flowers open simultaneously; flower (from base of 
ovary to apex of labellum) c. 4.5 cm long. Sterile bracts c. 13: 5.5-10 x 1.5- 
3.5 cm: soft (not rigid), spreading outwards from a loosely imbricating 
base: oblong-elliptic, strongly recurved: apex rounded, with subapical tooth 
c. 0.5 cm long: glabrous: pink with white margins. Outer floral bracts 
resembling the sterile bracts, c. 7.5 x 1-2 cm: inner floral bracts smaller c. 4 
x 0.7 cm. Bracteole 2-2.5 x c. 1.5 cm (when flattened): unequally 2-lobed. 
apex of lobe acute with c.l cm slit on opposite side: glabrous, translucent 
with a pink median band. Calyx 2.5-3 x c. 1.5 cm (when flattened); 3- 
lobed. each lobe with an apical tuft of short, brown hairs: tube with c.l. 5 
cm slit on opposite side: glabrous: white, deep pink at apex. Corolla tube 
1-1.5 cm long: inner surface densely covered with long hairs at entrance to 
tube: lobes (petals) longer than calyx. 2-2.5 cm long, apices rounded, slightly 
hooded, white with pink tips: dorsal lobe not hiding the anther and stigma. 
c. 0.7 cm wide, lateral lobes narrower, c. 0.4 cm. Labellum held erect: 
blade 1.8-2 cm long. c. 0.8 cm wide; deep red with a yellow margin except 
at basal lobes: margin of blade plain: apex rounded and retuse. Staminal 
tube 0.5-1 cm long above the base of the corolla-lobes: inner surface of 
tube sparsely covered with long hairs, more hairy below anther, (not warty). 
Filament short, c. 0.5 x 0.3 cm wide, adaxially hairy. Anther c. 0.7 x 0.3 cm; 
dark red, with white hairs on pollen sacs. Fpigynous glands 0.2-0.4 x 0.2- 
0.3 cm: comprising 2 rounded mounds, surrounding base of style, each 
mound deeply and irregularly lobed. apices glabrous, surface conspicuously 
warty. Ovary 0.4-0.6 x 0.3-0.5 cm; densely covered with appressed, white, 
short hairs; style white, hairy: stigma dark red, 0.3 cm across. Infructescence 
elongate, up to c. 19 cm but usually shorter x 10 cm in diam.: surmounted 
by withered flowers; withered, sterile bracts persistent until fruit maturity 
at base of fruit head: floral bracts persistent with fruits at their axils: number 
of fruits per head 15-25 (or more). Fruits obovoid; c. 2.5 x c. 3 cm wide at 
the top: densely covered with appressed short hairs; apex with remnant of 
calyx c. 3 cm long: pericarp smooth, not ridged longitudinally; pericarp 
0.2-0.3 cm thick: buff or pale green at maturity. Seeds irregularly angular, 
truncate: c. 0.4 x 0.3 cm wide. 



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Distribution: This species is widely cultivated and has been collected from 
several places in the Peninsula. Ridley (1924) reported the species wild in 
forests in Rawang. Selangor and in Tambun near Ipoh and the Taiping 
Hills in Perak. The species is described as native to the Peninsula. Java and 
Sumatra (Ridley, 1899). 

Notes: Locally known as kantan. this is commonly cultivated as a flavouring, 
the young inflorescences and fruits being used in curries. The newly opened 
inflorescence is beautiful but the bracts quickly turn black and rot. rendering 
it unsuitable for the cut-flower trade. 

One of the most conspicuous features of this species is the lengthening 
of the receptacle of the inflorescence during flowering, the spike thus 
forming an elongating cone within the involucral cup. 

Ridley (1899 p: 148 & 149) described the lip as white edged in 
Hornstedtia imperialis var. imperialis and the margins as yellow in var. 
speciosa. Ibrahim (1986) reported a form with white bracts and yellow 
flowers from Pulau Tioman. 

Specimens seen: PERAK: Tanjong Malim. Burkill & Haniff 14002 (SING): Ipoh. Ampang 
limestone cliffs. Burkill & Haniff 13934 (SING): Taiping. Batu Tegoh. Henderson 10139, 
10143, 10312 (SING): Larut Hills. King's Collector 3075 (SING): Telok Anson, Durian 
Sebatang. Haniff 15293 (SING): Taiping Waterfall, Ridley s.n. (SING): Bubu F.R.. Jalong. 
Symington SFN 39556 (KEP). SELANGOR: Rawang. Ridley 7811 (SING): Kanching. Bukit 
Takun, Stone 8787 (SING). JOHOR: Kota Tinggi. Ridley s.n. (SING). PAHANG: Pelangai 
or Manchis. Burkill & Haniff 16793 (SING); Ulu Tembeling. Sg. Sat. Henderson 22068 
(SING). TERENGGANU: Ulu Brang, Moysey & Kiah SFN 33873 (SING): Kuala 
Terengganu. Bukit Berangan. Sinclair & Kiah SFN 40891 (SING). 

2. Etlingera maingayi (Baker) R.M. Sm., Notes Roy. Bot. Gard. Edinb. 
43 (1986) 247; Lim. Folia malaysiana 1 (2000) 5. Syn.: Amomum maingayi 
Baker, in Hook. /., Fl. Brit. India 6 (1892) 235; Hornstedtia maingayi (Baker) 
Ridl.. in J. Str. Br. Roy. Asiat. Soc. 32 (1899) 150; Mat. Fl. Malay' Penins. 1 
(1907) 41; Phaeomeria maingayi (Baker) K. Schum.. Pflanzenr. Zingib. 20 
(1904) 266; Ridl.. Fl. Malay Penins. 4 (1924) 272; Holttum, Gard. Bull. 
Sing. 13 (1950) 180; Henderson. Malayan Wild Flowers. Monocots. (1954) 
155; Nicolaia maingayi (Baker) K. Larsen, Nat. Hist. Bull. Siam Soc. 23: 
574 (1970). Type: Malacca, Maingay 1586, s.v. (K). 

Figures 1A, 2A & B, 3A, 4A 

Rhizomes long, slender, creeping, c. 2.5 cm diam.; supported by stilt-roots 
10-30 cm long; scale-leaves overlapping: green flushed orange-pink. Leafy 
shoots 3-<\ m tall; 10-20 cm apart, forming clumps: young sheaths slightly 
longitudinally ribbed, with cross-bars covered by tufts of very short, white 



Etlingera iZingiberaceae) in Peninsular Malaysia 



209 



hairs; crushed sheaths with a pleasant, sour fragrance; ligule to 1.5 cm, 
apex entire or slightly bilobed, margin and surface with short, light-brown 
hairs; petiole to 0.5 cm, surface longitudinally ribbed. Laminas c. 17 pairs, 
(topmost) 25-62 x 3-5 cm, (middle) c. 66 x 10-12 cm, (basal) 20^3 x 5.5- 
8.5 cm; upper surface smooth, entirely green (young leaves sometimes 
brownish-pink all over on both surfaces or with brownish-pink bars on 
green lamina), lower surface glabrous, base cuneate, unequal. Peduncle up 
to c. 100 cm long, shortly horizontal, then erect, close to base of leafy 
shoot; sheaths c. 9, shortest at base 2-7 cm long, apex rounded, with 
subapical tooth c. 0.2 cm long, pink or purplish-red at apex, green towards 
base, not overlapping, widely spaced. Inflorescence raised well above the 
ground. Fertile apical part of inflorescence with well developed sterile bracts 
tightly imbricating, forming a firm, cup-like structure, 2-3 xc.4 cm; in side 
view the cup hiding c. 1/3 of the inflorescence head proper, the cup persistent 
until fruit maturity. Receptacle of inflorescence 1.5-2 cm long; apex inflated 
and conical; bearing c. 70 flowers; c. 9 flowers open simultaneously; flower 
(from base of ovary to apex of labellum) c. 4.5 cm long. Sterile bracts c. 8; 
3-4 x 4.5-5 cm; rigid, erect, subrotund, the whole concave towards the 
inside; apex broadly rounded, retuse, with a subapical tooth 0.2-0.5 cm 
long; outer surface with densely appressed, white hairs, red. Outer floral 
bracts like sterile bracts; c. 3.5 x 1.5-3 cm; inner floral bracts smaller, 3-3.5 
x 0.5-1.5 cm. Bracteole c. 2 x 1 cm (when flattened); apex 2-lobed and 
rounded with 0.5 cm slit on opposite side; densely, tawny hairy on entire 
surface. Calyx 2.5-3 x c.l cm (when flattened); 3-lobed, each lobe with a 
glabrous, subapical tooth 0.2 cm long; tube with c.l cm slit on opposite 
side; surface densely covered with appressed, tawny hairs; red. Corolla 
tube c. 1.5 cm long; inner surface densely covered with long hairs at entrance 
of tube; lobes (petals) shorter than calyx, c. 1.5 cm long, apices rounded, 
not hooded, red; dorsal lobe not hiding the anther and stigma, c. 0.5 cm 
wide, lateral lobes narrower, c. 0.2 cm. Labellum held erect; blade 1.5-2 
cm long, 0.6-0.8 cm wide; blade and basal lobes pink with a red median 
band not reaching the apex, margins of base white; margins of blade crisped; 
apex rounded and bilobed. Staminal tube c. 1 cm long above the base of 
the corolla-lobes; inner surface of tube densely covered with long hairs, 
not warty. Filament very short, c. 0.1 x 0.2 cm, adaxially hairy. Anther c. 0.8 
x 0.2 cm, red, hairy on pollen sacs. Epigynous glands c. 0.3 x 0.2 cm; 
comprising 2 rounded mounds, surrounding base of style, each mound 
minutely lobed, apices hairy, (not warty). Ovary c. 0.3 x 0.3 cm, glabrous; 
style red, hairy; stigma red, c. 0.2 cm wide. Infructescence spherical, small, 
5-7 x 5.5-7 cm in diameter; surmounted by withered flowers; sterile bracts 
persisting until fruit maturity at base of fruit-head; floral bracts persistent 
with fruits at their axils; 12 — 15 fruits per head. Fruits obovoid, c. 2.5 x 2.5- 



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3.5 cm wide at the top; glabrous; apex with remnant of calyx c. 3 cm long; 
pericarp smooth, not ridged longitudinally; pericarp 0.4-0.8 cm thick; bright 
red, shiny. Seeds irregularly angular, truncate, c. 0.4 x 0.4 cm. 

Distribution: In Peninsular Malaysia, this species had been collected from 
many localities in lowland forests. It is also found in Thailand (Smith. 
1986). 

Notes: This widely distributed species has involucral/sterile bracts of variable 
widths and apices. The inflorescence is recognizable from its red peduncle, 
small subglobose flower-heads and pink outer bracts entirely covered with 
appressed silvery-white hairs and rose-red flowers. 

Lim (2000) published a new variety, var. ovata (type at KEP), which 
has much broader leaves, 18.5 cm wide, compared with c. 12 cm wide and 
sessile, (not petiolate) in var. maingayi. Unfortunately, vegetative characters 
are less reliable than those of the inflorescence in Etlingera taxonomy. 
Within the same species, some leafy shoots may produce both petiolate 
and sessile leaves, as in E. pauciflora. Lim did not give any measurements 
of the inflorescence or its organs for a comparative study although his 
photographs showed specimens in flower. A more complete study is 
required for a taxonomic decision on the status of this variety. 

In the same publication. Lim (2000) rendered Holttum's var. 
longibracteata (type at SING) synonymous with var. maingayi. In his variety. 
Holttum (1950) described the involucral bracts as c. 3 x 1.5 cm (v. c. 3 x 2.5 
cm in var. maingayi), with the apex narrowly rounded instead of broadly 
rounded. Without stating sound reasons, Lim declared that var. 
longibracteata, "may be an aberration, and has so far not been found again 
near the type location - where the usual form is not uncommon". However, 
not being able to relocate a taxon at the type-site (moreover, Tembeling' 
covers a large area) is insufficient grounds for eliminating Holttum's var. 
longibracteata. The variety should be maintained until detailed comparative 
studies with var. maingayi are made. 

Specimens seen: PERLIS: Lubuk Sireh, Hutan Simpan Mata Ayer, Wan Fadhilah HI 931 
(KLU). KEDAH: Pass to Kroh from Baling, Comer SFN 31569 (SING); Yan, G. Jerai, 
Batu Hampar. Saw FRI 44640 (KEP). PERAK: Upper Perak, Wray s.n. (SING); Temengor. 
Ridley 14419 (SING). SELANGOR: Dusun Tua. Ridley 7801 (SING). JOHOR: Sedili 
River, below Mawai, Corner SFN 36969 (2 sheets) (KEP, SING,); Batu Pahat. Ridley s.n. 
(SING); Ulu Kahang, Holttum 10902 (SING); Kota Tinggi, G. Panti F.R.. West. Saw FRI 
37746 (KEP). KELANTAN: Kuala Lebir, Gimlette s.n. (SING); Gua Musang. Saw FRI 
37403 (KEP). PAHANG: Kota Glanggi, Furtado s.n. (SING); Ulu Sg. Sat, Mohd. Shah & 
Mohd. Noor MS 1772 (KEP. SING); Genting Highlands, Awana Trail, Saw FRI 37406 
(KEP); Taman Negara, Sg. Relau, Chua, Mustapa & Apok FRI 40607 (KEP). 



Ellingera < Zingiberaceae ) in Peninsular Malaysia 



211 



3. Etlingera fulgens (Ridl.) C.K. Lim. Folia malaysiana 1 (2000) 6. Syn.: 
Hornstedtia fiilgens Ridl., J. Sir. Br. Roy. Asiat. Soc. 32 (1899) 149; Mat. 
Fl. Malay. Penins. (1907) 40; Phaeomeria fulgens (Ridl.) K. Schum., 
Pflanzenr. Zingib. (1904) 262: Ridl., Fl. Malay Penins. 4 (1924) 272; Holttum, 
Gard. Bull. Sing. 13 (1950) 180; Nicolaia fiilgens (Ridl.) K. Larsen, Nat. 
Hist. Bull. Siam Soc. 23 (1970) 574. Type: Perak, Ridley s.n. (SING, iso). 

Figures 1C, 2E & F, 3B, 4C 

Rhizomes just below ground; c. 3 cm diam.; scale-leaves overlapping, 
creamy-white. Leafy shoots 4-5 m tall; 10-14 cm apart, forming clumps; 
young sheaths faintly ribbed longitudinally, with cross-bars not covered by 
tufts of short, white hairs; crushed sheaths with a pleasant, sour fragrance; 
ligide 1.5-2 cm. apex broad, entire or slightly bilobed, margin with short, 
light brown hairs; petiole 1.5-2 cm, surface longitudinally ribbed. Laminas 
c. 15 pairs; (topmost) c. 31 x 4 cm, (middle) 64-96 x 14-19 cm, (basal) 12- 
28 x 4.5-8.5 cm: upper surface smooth, glossy, dark green; lower surface 
glabrous; young leaves with lower surface conspicuously dark purple-red, 
older leaves with only petiole and midrib purple-red; base variable, blunt 
or rounded, unequal. Peduncle up to c. 100 cm, shortly horizontal at first, 
then erect, aerial, close to base of leafy shoot; sheaths c. 12. shortest at 
base 2.5-12 cm long, apex rounded with subapical tooth 0.1 cm long, green, 
not overlapping, widely spaced. Inflorescence raised well above the ground. 
Fertile apical part of inflorescence with well developed sterile bracts tightly 
imbricating, forming a firm, broad cup-like structure c. 8.5 x c. 7.5 cm, in 
side view the cup almost entirely hiding the inflorescence head proper, the 
cup not persisting until fruit maturity. Receptacle of inflorescence c. 1.5 cm 
long, apex inflated and rounded: bearing c. 60 flowers; 3-6 flowers open 
simultaneously; flower (from base of ovary to apex of labellum) c. 5 cm 
long. Sterile bracts c. 7; 5-7 x 3.5-5.5 cm; rigid, erect, subrotund, strongly 
incurved at apex: apex broadly rounded, slightly retuse, with subapical 
tooth hardly 0.1 cm long: outer surface glabrous to sparsely hairy all over 
with white hairs, deep pink with green margins. Outer floral bracts 
resembling sterile bracts, 4-5 x 2-3.5 cm; inner floral bracts smaller, 3-4 x 
0.5-1.5 cm. Bracteole 2.5 x c. 1 cm (when flattened); 2-lobed, apex of lobe 
acute with 1.5 cm slit on opposite side; glabrous, translucent, tinged red. 
Calyx c. 4 x 1 cm (when flattened): 3-lobed, each lobe with a glabrous 
subapical tooth less than 0.1 cm: tube with c. 2 cm slit on opposite side, 
glabrous, red. Corolla-tube c. 1.5-2 cm long, inner surface densely covered 
with long hairs at entrance into tube: lobes (petals) longer than calyx, 2 cm 
long, apices rounded, slightly hooded, red with white tips; dorsal lobe not 
hiding the anther and stigma, c. 0.5 cm wide, lateral lobes c. 0.3 cm. Labellum 



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held erect; blade 1.8-2 cm long, c. 0.8 cm wide; blade and basal lobes pink- 
red with a yellow margin except at basal lobes; margins of blade plain, 
apex rounded and retuse. Staminal tube c. 0.5 cm long above the base of 
the corolla-lobes; inner surface of tube sparsely covered with long hairs, 
more hairy below the anther, (not warty). Filament short, c. 0.3 x 0.2 cm, 
adaxially hairy. Anther c. 0.8 x 0.4 cm, pink-red, hairy on pollen sacs. 
Epigynous glands 0.5-0.6 x 0.2 cm; comprising 2 rounded mounds, 
surrounding base of style, each mound deeply and irregularly lobed, apices 
hairy, surface conspicuously warty. Ovary c. 0.6 x 0.3 cm; densely appressed 
hairy with white, short hairs; style red, hairy; stigma pink-red, c. 0.3 cm 
across. Infrnctescence globular, flat-topped, large; c. 11 x 13 cm in diameter; 
not surmounted by withered flowers; sterile and floral bracts completely 
disintegrated at fruit maturity; 30 — 35 fruits per head. Fruits ovoid; c. 4-5 x 
2.5-3.5 cm at the middle; densely covered by appressed short hairs; apex 
with remnant of calyx 3-4 cm long; pericarp smooth, (not ridged 
longitudinally); pericarp 0.2-0.4 cm thick; pale green at maturity. Seeds 
irregularly angular, truncate; c. 0.4 x 0.3 cm. 

Distribution: In Peninsular Malaysia, this species is not common. Holttum 
(1950) reported it was in cultivation in Singapore. Larsen (1970) recorded 
it occurs in southern Thailand. 

Notes: This species is recognized by its shiny leaves that are dark green 
above and when young are conspicuously purple-red beneath but turn 
green when old and have wavy margins, and by its inflorescences with 
rounded, sterile bracts, strongly incurved at the apex, rose-red with greenish 
white margins, forming a shallow, wide cup. The young plants sometimes 
produce short stilt-roots but these are not observed in adult plants. 

Smith (1986) had combined this taxon with a Javan one as Etlingera 
hemisphaerica (Blume) R.M. Sm.. Lim (2000) rejected this decision and 
resurrected the epithet fulgens for the Peninsular Malaysian taxon. Among 
the arguments he presented were that Smith (1986), in making the 
combination E. hemisphaerica, did so without having viewed the types of 
either Ridley's Malayan Hornstedtia fulgens or Blume's Javan Elettaria 
hemisphaerica. However, Lim did not make clear whether he had actually 
seen the types of these two species himself nor did he give a critical 
assessment of the inflorescence characters of both the species. The 
arguments presented by Lim for his decision are therefore not conclusive. 
Since I have not seen Blume's type from East Java, and to avoid any 
unnecessary changes until the types of both species are examined, E. fulgens, 
is used in the present work for the taxon from Peninsular Malaysia. 



Etlingera I Zingiberaceae ') in Peninsular Malaysia 



213 



Specimens seen: PERAK: Jor. Batang Padang. Henderson 10854 (SING). PAHANG: Krau 
Wildlife Reserve. Hulu Sg. Lompat. Chita FR I 41742 (KEP). TERENGGANU: Ulu Brana. 
Moysey & Kiah S FN 33670 (SING). 

4. Etlingera venusta (Ridl.) R.M. Sm.. Notes Roy. Bot. Gard. Edinb. 43 
(1986) 250. Syn.: Hornsiedtia venusta Ridl.. in J. Str. Br. Roy. Asiat. Soc. 
32 (1899) 149: Mat. Fl. Malay Penins. (1907) 40: Phaeomeria venusta (Ridl.) 
K. Schum.. Pflanzenr. Zingib. (1904) 264: Ridl.. Fl. Malay Penins. 4 (1924) 
272: Holttum. Gard. Bull. Sing. 13 (1950) 182: Nicolaia venusta (Ridl.) K. 
Larsen. Nat. Hist. Bull. Siam Soc. 23 (1970) 575. Type: Selangor. Genting 
Bidai. Ridley 7810 (K. holo: SING. iso). 

Figure 5 

Rhizomes just below ground: 2.5^1.5 cm diam.: scale-leaves overlapping, 
green. Leafy shoots 2.5-4 m tall: 10-18 cm apart, forming clumps: young 
sheaths longitudinally ribbed, with cross-bars not covered by tufts of short, 
white hairs: crushed sheaths with a pleasant, sour fragrance: ligule 2-3 cm: 
apex deeply bilobed: margin and surface densely covered with short, brown 
hairs: thin, papery, upper portion breaking off easily: petiole inconspicuous, 
always subsessile. Laminas c. 10 pairs: (topmost) 51-64 x 9.5-13 cm. (middle) 
66-80 x 14-20 cm. (basal) 45-54 x 13-14.5 cm: lamina prominently ribbed: 
entirely green on both surfaces: lower surface glabrous: base cuneate always. 
Peduncle to 60-80 cm: shortly horizontal, then erect, aerial, close to base 
of leafy shoot: sheaths c. 12. shortest at base 2-9 cm long, apex rounded 
with subapical tooth 0.2 cm long, flushed red at apex, green towards base, 
not overlapping, widely spaced. In florescence raised well above the ground. 
Fertile apical part of in florescence with well developed sterile bracts, tightly 
imbricating, forming a firm, deep cup-like structure 8-9 x c. 7.5 cm: in side 
view the cup almost entirely hiding the inflorescence head proper: cup not 
persisting until fruit maturity. Receptacle of inflorescence 2-3 cm long: 
apex inflated and conical: bearing 60-70 flowers: 1-5 flowers open 
simultaneously: flower (from base of ovary to apex of labellum) 5-5.3 cm 
long. Sterile bracts c. 8: 7.5-12 x 4-5.5 cm: rigid, erect, oblong-elliptic, 
strongly recurved at apex: apex acute, with subapical tooth c. 0.5 cm: outer 
surface glabrous or sparsely covered with white hairs: bracts entirely purple- 
pink or with some white towards apex, pale green at base. Outer floral 
bracts resembling inner sterile bracts but narrower, 7.5-8 x 2.5-3.7 cm; 
inner floral bracts smaller, 2-6 x 0.5-1.5 cm. Bracteole c. 3 x 1.5 cm (when 
flattened): 2-lobed. apex of lobe acute: with c. 2 cm slit on opposite side: 
glabrous, translucent, pink towards apex. Calyx 4-4.3 x c. 1.5 cm (when 
flattened): 2- or 3-lobed. each lobe with a glabrous or hairy subapical tooth 
0.1 cm: tube with 2-2.5 cm slit on opposite side: glabrous, with brown hairs 



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Figure 5. Etlingera venusta. A. base of peduncle; B, inflorescence; C, sterile bract; D, floral 
bract; E, flower with bracteole, side view; F, bracteole; G, calyx; H, flower dissected with style 
displaced; I, infructescence; (A from dry material, B-I from spirit material of Khaw KSH 
244). 



Etlingera (Zingiberaeeae) in Peninsular Malaysia 



215 




216 



Gard. Bull. Singapore S3 (2001) 



at apex only; white, deep purple-red towards apex. Corolla-tube t— 1.2 cm 
long, inner surface with long hairs at entrance into tube; lobes (petals) 
shorter than calyx, c. 1.5 cm long, apices rounded, slightly hooded, white 
with pink tips; the dorsal lobe not hiding the anther and stigma, 0.5-0.7 cm 
wide, lateral lobes 0.4-0.5 cm. Labellum held erect; blade narrowed to 
base without widening to form basal lobes; 2-2.2 x 1.2-1.3 cm; white with a 
red median band not reaching the apex, without red lines radiating from 
the red band towards the margin; margin slightly crisped or plain; apex 
rounded, (not retuse). Staminal tube 1.2-1.3 cm long above the base of the 
corolla-lobes; inner surface of tube with long hairs, (not warty). Filament 
very short, c. 0.1 x 0.2-0.3 cm wide, adaxially hairy. Anther c. 1 x 0.3 cm, 
pale yellow or white, with white hairs on pollen sacs. Epigynous glands c. 
0.5 x 0.2 cm; comprising 2 rounded mounds, surrounding base of style, 
each mound deeply and irregularly lobed, apices hairy, surface warty 
towards the apices. Ovary 0.4-0.7 x 0.4-0.7 cm, glabrous; style white, hairy; 
stigma pale pink, 0.3 cm across. Infructescence globular; large, c. 9 x 13-15 
cm diam.; not surmounted by withered flowers; sterile and floral bracts 
completely disintegrated at fruit maturity; 15 — 20 fruits per head. Fruits 
ovoid, c. 5 x c. 3.5 cm wide at the middle; glabrous, apex with remnant of 
calyx 3-4.5 cm long; pericarp smooth, not ridged longitudinally; pericarp 
0.4-0.5 cm thick; bright red or pink, shiny. Seeds irregularly angular, 
truncate; c. 0.4 x 0.3-0.5 cm wide, widest at apex. 

Distribution: In Peninsular Malaysia, this species is collected mainly from 
Perak, Selangor and Pahang. As it has not been recorded elsewhere, P. 
venusta is therefore endemic to Peninsular Malaysia. 

Notes: This species is distinguished from E. corneri by the absence of stilt- 
roots, its subsessile, broader leaves with cuneate bases; its rose-pink 
inflorescences formed by bracts with acute apices and infructescences of 
red fruits not surrounded by bracts, which completely decay during fruit 
development. 

Specimens seen: PERAK: Ulu Batang Padang, Rid!. 13835 (SING); Tapah Hills. Sg. Woh, 
Ng FRI 1330 (KEP, SING); Tapah Hills, S. Keteh, Whitmore FRI 15668 (KEP); Hulu 
Perak, Temenggor F.R., Sg. Samlor. Khaw KSH 378 (KEP). PAHANG: Genting Highlands, 
Awana Hotel Trail. Saw FRI 36448 (KEP). 

5. Etlingera corneri J. Mood & H. Ibrahim, Nord. J. Bot. 20 (24. 8. 2000) 
278-283. Syn.: Etlingera terengannuensis C.K. Lim, Folia malaysiana 1 (2000) 
9. Type: Thailand, Narathiwat Province, 1991, Vitoon, Lyon 9L 433, s.v. 
(KLU, SING). 



Ellingera f Zingiberaceae) in Peninsular Malaysia 



217 



Figure 6 

Rhizome above leaf-litter or just below soil level. 3-4.5 cm diam.: supported 
by pinkish-orange stilt-roots. 10-25 cm long: scale-leaves overlapping, green 
tinged dull red. Leafy shoots 2.5-4 m tall: close together. 6-14 cm apart, 
forming clumps: sheaths longitudinally ribbed, with cross-bars not covered 
by tufts of short, white hairs, when crushed emitting a pleasant, sour 
fragrance: ligule 1.5-2 cm, apex deeply bilobed, margin and surface densely 
covered with short hairs: petiole 1.5-4 cm, surface reticulate. Laminas 10- 
11 pairs: (topmost) 36-47 x 5.5-8 cm, (middle) 57-65 x 13-16 cm, (basal) 
14-35 x 5-10 cm: apex shortly-pointed, c.l cm long: base variable, mostly 
rounded, occasionally unequal, sometimes cuneate: entirely green on both 
surfaces when young: upper surface prominently ribbed: lower surface 
glabrous: margin red. crisped, with short white or light brown hairs. Peduncle 
c. 90 cm, shortly horizontal to c. 4 cm, then erect, close to base of leafy 
shoot: sheaths c. 12, shortest at base, 2.5-13 cm long, apex bilobed with 
subapical tooth to c. 0.5 cm long, surface with short hairs, red at apex, pale 
green towards base; not overlapping, widely spaced. Inflorescence raised 
well above the ground. Fertile apical part of inflorescence with well 
developed sterile bracts tightly imbricating, forming a firm. deep, cup-like 
structure 8-9 x c. 7.5 cm: the cup almost entirely hiding the inflorescence- 
head proper in side view: the cup persistent until fruit maturity. Receptacle 
of inflorescence 1-1.5 cm long; apex inflated, conical or rounded: bearing 
50-60 flowers: 1-5 flowers open simultaneously: flower (from base of ovary 
to apex of labellum) 5.3-6 cm long. Sterile bracts 8-14; 6.5-8.5 x 2-5.5 cm; 
rigid, erect, oblong-elliptic, recurved at apex; apex subtruncate, retuse, 
with subapical tooth c. 0.5 cm long: outer surface of bract glabrous, densely 
covered with appressed white hairs at base only: bracts bright red. white or 
pale green at base. Outer floral bracts resembling the sterile bracts but 
smaller. 5.5-7.5 x 2-4 cm; inner floral bracts smaller still. 3.5-4 x 0.5-1.5 
cm. Bracteole tubular at base; c. 3.5 x 1.5 cm when flattened: unequally 2- 
lobed. apex of lobe acute or rounded with c. 2 cm slit on opposite side; 
surface with white hairs; white with red apex. Calyx tubular at base; 4-4.4 
x c. 1.5 cm when flattened; 3-lobed, each lobe with a glabrous or hairy 
subapical tooth c. 0.1 cm long; tube with c. 2.5 cm slit on opposite side; 
glabrous, with appressed white hairs at base only; white, red at apex. 
Corolla-tube 1.5-2 cm long, inner surface densely covered with long hairs 
at entrance into tube; lobes (petals) shorter than calyx, c. 2 cm long, apices 
rounded, not hooded, pink; dorsal lobe not hiding the anther and stigma. 
c. 0.5 cm wide, lateral lobes narrower, c. 0.4 cm wide. Labellum held erect, 
blade narrowed to base without widening to form basal lobes; c. 2 x 1.2-1.3 
cm; white with a red median band not reaching the apex, with red lines 



218 



Card. Bull. Singapore 53 (2001) 




1 cm 

1 cm 




Figure 6. Etlingera cortieri. A. leaf base and tip: B, inflorescence; C. sterile bract, adaxial 
view: D, sterile bract, lateral view; E. floral bract, adaxial view; F. flower with bracteole, lateral 
view; G, flower with bracteole. adaxial view; H. bracteole. flattened, adaxial view; I. calyx, 
flattened, adaxial view; J, flower dissected with style displaced; K, epigynous glands, adaxial 
view; L, epigynous glands, abaxial view; M, infructescence with persistent involucral and floral 
bracts displaced; (A. B from fresh material, C-L, spirit, M. dry; from Khaw KSH 601 ). 



220 



Card. Bull. Singapore 53 (2001) 



radiating from the red band towards the margin; margin slightly crisped; 
apex rounded, slightly retuse. Staminal tube 0.9-1 cm long above the base 
of the corolla-lobes, inner surface of tube densely covered with long hairs, 
(not warty). Filament very short, c. 0.1 x 0.2-0.3 cm, adaxially hairy. Anther 
c. 1 x 0.3 cm, red at apex, white at base, with yellow hairs on pollen sacs. 
Epigynous glands c. 0.4 x 0.2 cm; comprising two rounded mounds, 
surrounding base of style, each mound deeply and irregularly lobed, apices 
glabrous, surface warty towards the apices. Ovary 0.4-0.6 x c. 0.4 cm, 
glabrous, white; style white, hairy; stigma pink, c. 0.2 cm across. 
Infructescence globular, large, c. 9 x 10-11 cm, not surmounted by withered 
flowers, sterile bracts surrounding fruits persisting to fruit maturity, 8 — 17 
fruits per fruit head. Fruit ovoid; c. 4 x 2.5 cm at the middle; glabrous; apex 
with remnant of calyx 3-4.5 cm long; bright red or pink, shiny; pericarp 
smooth, not ridged longitudinally, 0.3-0.4 cm thick. Seeds 35-45 in each 
fruit; covered with a white, pulpy aril; dark brown; irregularly angular 
truncate; c. 0.4 x 0.3-0.5 cm, widest at apex. 

Distribution: This attractive species of horticultural potential has been 
recorded in the Peninsula from the states of Kedah, Perak, Pahang and 
Terengganu. It is not common and there are few collections. It is also 
found in southern Thailand (Mood & Ibrahim, 2000). 

Notes: The confusion over Etlingera corneri and E. venusta is discussed in 
detail by Mood & Ibrahim (2000). E. corneri is distinguished from E. 
venusta by its stilt roots, petiolate leaves usually with rounded bases, its 
bright red inflorescence in the form of a deep cup; the outer bracts that are 
strongly recurved and have subtruncate apices, and the infructescence of 
red fruits surrounded by persistent bracts. 

The plants from northern Perak, on which the above description is 
based, have leaves that are entirely green on both surfaces, even when 
young. This character differs from that described by Mood & Ibrahim 
(2000): "dark green above, ... green (or purple) below ...". For two specimens 
from Johor, which were previously annotated as Phaeomeria venusta and 
which were cited by Mood & Ibrahim as being E. corneri, the field notes 
read: "leaves dark green, glossy with undulate edge, purple beneath, though 
old leaves may lose the purple" (Corner SFN 32778, KEP, SING) and 
"dark green above, purple beneath, glossy on both surfaces" (Sinclair 10596, 
SING). This leaf character is reminiscent of E. fulgens. As I am uncertain 
of the identity of Corner SFN 32778 and Sinclair SFN 10596, 1 have excluded 
them from the list below of specimens seen. The field notes in Corner SFN 
32778 also mention "stilts 2-4" high", shorter than those described here. 
Until Mood and Ibrahim (2000) clarified the confusion over the identities 



Etlingera ( Zingiberaceae) in Peninsular Malaysia 



221 



of E. venusta and E. corneri, these two species were often misidentified. 
The photographs in Weber (1995) showing the inflorescence and 
infructescences of E. venusta from Sekayu, Terengganu are actually those 
of E. corneri. This was noted by Mood and Ibrahim (2000). Similarly, 
photographs of E. venusta infructescences in Larsen et al. (1999) are in fact 
those of E. corneri. while the photograph showing "clusters of horn-like 
Etlingera fruits" are the fruits of E. venusta. 

Specimens seen: KEDAH: Ulu Muda F.R., Baling. Waduk, K.H. Lim, LKH 158 (KEP). 
PERAK: Hulu Perak.Temenggor F.R.. Pos Chiong. Sg. Sara, Khaw KSH 373 (KEP); Sg. 
Tiang. Khaw KSH 586, 601, 606 (KEP). PAHANG: Kuantan, Sg. Pohoi. Mohd. Shah. 
Sidek & Samsuri MS 3787 (KEP). TERENGGANU: Ulu Brang. Moysey & Kiah SFN 
33721 (KEP. SING): Hulu Terengganu F.R.. Hutan Lipur Sekayu. Anthonysamy SA 648 
(KEP): Ulu Terengganu (Extension) F.R.. Cpt. 43, Saw FRI 39880 (KEP); Ulu Terengganu 
F.R.. C.K Lim LIM 4229 (KEP). 

6. Etlingera pieeae Khaw. sp. nov. 

Etlingera triorgyali affinis sed inflorescentia supra terram elevata. pedunculo 
saltern duplo longiore, bracteis involucralibus tantum basin luxe imbricatis 
ergo cupulam involucralem conspicuam non formanti, inflorescentia inter 
bracteas a latere visibili, bracteis rigidis leviter expansis, oblanceolatis. 
longitudinaliter inflexis non recurvatis, pallide viridis marginibus et apicibus 
rubris et longitudinaliter rubro-striatis, apice inflato axis florifero conico, 
flore, bractea involucrale, bracteis. bracteolis et calycibus brevioribus et 
angustioribus, corolla calyce longiore loba dorsali antheram et stigma 
occultanti, tubo corollae nutlto breviore, labello erecto loba media breviore 
et angustiore, tubo staminali breviore, infructescentia minore vestigiis 
bractearum involucrarum persistentibus, fructibus paucis minoribus, 
pericarpio laeve non porcato recedit. Typus: Perak. Hulu Perak. Belum 
F.R., Sg. Beruar, Khaw KSH 420 (holotypus KEP; isotypus KLU). 

Figure 7 

Rhizomes just below soil level; 3^ cm diam.: scale-leaves overlapping, 
green. Leafy shoots to 4-5 m tall; 7-18 cm apart, forming clumps; sheaths 
conspicuously longitudinally ribbed with cross-bars covered by short, white 
hairs: when crushed with a strong, peppery scent: ligule 2-2.5 cm long, 
apex acute, margin and surface densely covered with short, brown hairs; 
petiole 2.5-4 cm long, surface reticulate. Laminas 13-14 pairs; (topmost) 
68-73 x 7-9 cm. (middle) 87-93 x 16-21 cm, (basal) c. 55 x 13 cm; when 
young entirely green on both surfaces: upper surface prominently ribbed: 
lower surface and margin with soft, short hairs; base cuneate, unequal; 



222 



Card. Bull. Singapore 53 (2001 ) 



1 cm 

1 cm 




Figure 7. Etlingera pieeae S.H. Khaw, sp. nov. A, young inflorescence; B, inflorescence with 
sheaths and flower-head; C, sterile bract, abaxial view; D, sterile bract, adaxial view; E. floral 
bract, abaxial view; F, bracteole, flattened, abaxial view; G, calyx, flattened, abaxial view; H, 
flower dissected with style displaced; I, stigma, abaxial view; J, stigma, lateral view; K. epigynous 
glands, adaxial view; L. epigynous glands, abaxial view; M, infructescence with fruits surrounded 
by remnants of sheaths and bracts with skeletonized veins. (A-L from spirit material of Khaw 
KSH 420, M from dry material.) 



224 



Card. Bull. Singapore 53 (2001 > 



apex shortly pointed, c. 1 cm long. Peduncle to c. 48 cm long, horizontal, 
then erect, aerial, close to base of leafy shoot; sheaths c. 12, shortest at 
base from 1.5-13 cm, apex rounded with a sharp, hard, subapical spine c. 
0.1 cm, overlapping, green, striped deep pink, dull purple-red at apex. 
Inflorescence raised well above the ground. Fertile apical part of inflorescence 
with well developed sterile bracts, loosely imbricating at the bases only, 
(not forming any conspicuous sub-cylindric structure), in side view the 
inflorescence head proper visible between the bracts. Receptacle of 
inflorescence c. 1 cm long, apex inflated and conical; bearing 20-35 flowers; 
7-12 flowers open simultaneously; flower (from base of ovary to apex of 
labellum) c. 9 cm long long. Sterile bracts 5-8; 9-12 x 1.5-3 cm; rigid, 
spreading slightly, oblanceolate, longitudinally inflexed but not recurved; 
apex rounded, retuse, crisped, with subapical hairy spine c. 0.2 cm long; 
outer surface with white hairs, densely appressed hairy at apex and base; 
surface pale green, apex and margin red and longitudinally striped reddish- 
pink. Outer floral bracts like inner sterile bracts; 8.5-9.5 x 1.5-2.5 cm; inner 
floral bracts smaller, c. 6 x 0.5-1 cm. Bracteole 5-6.5 x c. 1.5 cm (when 
flattened); 2-lobed, apex of each lobe rounded with c. 3 cm slit on opposite 
side; surface densely covered with tawny hairs; pink. Calyx 4.5-6 x c. 1.5 
cm (when flattened); 3-lobed, each with a glabrous, subapical tooth c. 0.1 
cm; tube with c. 2 cm slit on opposite side; surface densely covered with 
appressed white hairs. Corolla tube c. 4 cm long, inner surface densely 
covered with long hairs at entrance of tube, rest of tube glabrous; lobes 
{petals) 1-2 cm longer than calyx, c. 2 cm long, apices rounded, not hooded, 
margins crinkled, red; dorsal lobe hiding the anther and stigma, c. 1 cm 
wide, lateral lobes narrower, c. 0.5 cm. Labellum held erect, blade elongate, 
3.5-4 cm long, c. 2 cm wide; blade and basal lobes entirely bright pink-red; 
margin of blade crisped and crinkled; apex broadly rounded, slightly retuse. 
Staminal tube c. 0.5 cm long above the base of the corolla-lobes; inner 
surface of tube densely covered with long hairs especially below the anther, 
rest of surface densely covered by warts (glands?). Filament short, c. 0.3 x 
0.4 cm, adaxially hairy. Anther c. 1 x 0.3 cm, white, hairy on pollen sacs. 
Epigynous glands 0.5-0.6 x c. 0.2 cm; comprising 2 fleshy, compressed 
blade-like structures, not encircling base of style, each blade not further 
lobed, apex glabrous, shortly-pointed, (surface not warty). Ovary c. 0.5 x 
0.4 cm, densely covered with appressed, white, short hairs; style white or 
pale pink, hairy; stigma pink, c. 0.4 cm across. Infructescence a small spherical 
head, c. 2.5 x 5 cm (when dry), not surmounted by withered flowers, 
surrounded by remnants of sterile bracts with skeletonized veins persistent 
at fruit maturity, c. 10 fruits per fruit head. Fruit obovoid; c. 1.5 x 1 cm at 
the top (when dry); buff coloured; densely covered with appressed, short 
hairs; apex with remnant of calyx base only, c. 0.5 cm long (when dry); 



Etlingera (Zingiberaceae) in Peninsular Malaysia 



225 



pericarp smooth, not ridged longitudinally; pericarp 0.1-0.2 cm thick. Seeds 
irregularly angular, truncate. 

Distribution: Endemic to Peninsular Malaysia: Perak and Kedah (in Ulu 
Muda F.R.. Baling. Waduk. sighted only, not collected). 

Ecology: By banks of shaded as well as more open, unshaded, small streams 
in lowland forest at c. 300 m altitude. 

Notes: Etlingera pieeae closely resembles E. triorgyalis in vegetative 
characters. Both have tall leafy-shoots with sheaths that emit a strong 
peppery scent when crushed. They are often found growing near each 
other. However, its inflorescence shows that E. pieeae is a distinct species. 
It has a combination of floral characters of both the Phaeomeria and 
Achasma groups of Etlingera. It resembles the Phaeomeria group in having 
a long peduncle with its flower-head held well above the ground, but has 
overlapping peduncle-sheaths and a long labellum like those of the Achasma 
group. Its flower-head is neither cup-shaped (typical of the Phaeomeria 
group) nor sub-cylindric (most of the Achasma inflorescences). Instead, its 
inflorescence with slightly spreading oblanceolate. sterile bracts, loosely 
imbricating at the bases only, resembles that of E. pauciflora of the Achasma 
group. This combination of features in E. pieeae from the two groups 
makes it an interesting species, lending support to the union of Phaeomeria 
and Achasma into a single genus, Etlingera. 

Table 2 summarises the differences between E. pieeae and E. 
triorgyalis. its closest relative. 

This species is named for Mr. Abdullah Piee, who first discovered 
this new species, in appreciation for his assistance and companionship in 
ginger field trips. 

Specimens seen: PERAK: Hulu Perak. Temenggor F.R.. Sg. Emban. Khaw KSH 603 (KLU), 
KSH 604 (KEP): Belum F.R., Sg. Beruar. Khaw KSH 420 (KEP. KLU). 

7. Etlingera pauciflora (Ridl.) R.M. Sm., Notes Roy. Bot. Gard. Edinb. 
43 (1986) 248. Syn.: Hornstedtia pauciflora Ridl.. in J. Str. Br. Roy. Asiat. 
Soc. 32 (1899) 144; Mat. Fl. Malay. Penins. (1907) 38; Fl. Malay Penins. 4 
(1924) 270: Achasma pauciflorum (Ridl.) Holttum, Gard. Bull. Sing. 13 
(1950) 187. Type: Selangor, Gua Batu, Ridley 8174, s.v. (K). 

Figure IE, 21 & J, 3H, 4 E-G 

Rhizomes 6-13 cm below the ground; c. 0.6 cm diam.; scale leaves 
overlapping, white. Leafy shoots 2-4 m tall; close together or widely spaced. 



226 Card. Bull. Singapore 53 (2001 ) 



Table 2. Characters that distinguish Etlingera pieeae from E. triorgyalis. 







Li,, it luf £ yciiics 


Peduncle length (cm) 


to c. 48 


5-15 


Inflorescence 


raised well above the 


subterranean except for 




ground 


apices of sterile bracts 


flower-head 


sterile bracts loosely 


and flowers 




imbricating at the bases 


sterile bracts tightly 




only, not forming any 


imbricating, forming a 




conspicuous suDcynnuer 


narrow subcylinder. 




the flower head visible 


the subcylinder hiding 




between the bracts in side 


r 7/3 of the flower-head 




V 1C w 


in ciHp» \ri(*\\i 
111 MUC V1CW 


niimhpr r\f fln\x/f*r*; 
11UU1UC1 Ul 11UWC1 J 


ZU— JD 


tU-JU 


flower length (cm) 


c. 9 


c. 13.5 


sterile bracts 






form 


rigid, spreading slightly. 


rigid, erect, oblong- 




oblanceolate. 


obovate. recurved at 




lnnoitnrlinnllv in fl p ypH 

1 * ' 1 1 1 l 11 LI 1 1 1 l l l 1 \ M ! 1 1 L \ L U 






but not recurved 




width (cm) 


1.5-3 


3-6 


colour 


pale green, apex and 
margin red, longitudinally 
striped red 


white or pale green, 
deep pink towards apex 




corolla lobes 


1-2 cm longer than calyx 


LUUill IU L,CllVA 




dorsal lobe hiding anther 


rlor^n 1 lohp not huiino 




and stigma 


Cllllll^l Ml IU 'II — 1 1 1 <. 1 


tube length (cm) 


c. 4 


5 X-fi 5 


labellum length (cm) 3.5-4 


c. 5 


staminal tube 


c. 0.5 


c. 1.2 


length (cm) 






Infructescence size (cm) 


c. 2.5 x 5 (dry) 


c. 6 x 10 


persistence of 


the bracts partially decay. 


bracts completely 


sterile bracts at 


remnants of bracts with 


decayed 


fruit maturity 


skeletonized veins 




surrounding infructescence 




number of fruits 


c. 10 


20-35 


Fruit 






colour 


buff 


dark red or pink 


size (cm) 


c. 1.5 x 1 (dry) 


c. 3x3 cm 


pericarp 


smooth, not ridged 


ridged longitudinally. 




longitudinally 


each ridge with many 
irregular fine spine- 
like processes 



Ellingera (Zingiberaceae) in Peninsular Malaysia 



227 



9-80 cm apart; sheaths slightly longitudinally ribbed with cross-bars covered 
by tufts of short, white hairs (clearer in young sheaths), without distinct 
scent when crushed: ligule 1-2.5 cm; apex rounded, entire; margin with 
short, brown hairs; surface glabrous: lamina often sessile or with petiole 
0.5-1 cm long in laminas with unequal bases, surface reticulate. Laminas c. 
11 pairs: (topmost) c. 37 x 6.5 cm, (middle) 44-65 x 9-11 cm, (basal) c. 40 x 
12.5 cm; upper surface prominently ribbed; entirely green on both surfaces 
when young; lower surface glabrous: margins crisped, red with brown hairs; 
base cuneate, sometimes unequal. Peduncle 5-10 cm long; horizontal, then 
erect, entirely subterranean at base of leafy shoot or some distance away: 
sheaths c. 8. shortest at base 0.5-5.5 cm, apex acute with stiff apical spine c. 
0.1 cm, overlapping, white. Inflorescence entirely subterranean except for 
apices of sterile bracts and flowers. Fertile apical part of inflorescence with 

1- 3 well developed sterile bracts loosely imbricating at the bases only, not 
forming any conspicuous, sub-cylindric structure; in side view the 
inflorescence head proper visible between the bracts; the sterile bracts not 
persisting to fruit maturity. Receptacle of inflorescence c. 0.5 cm long; apex 
inflated and rounded, bearing 1-3 flowers; 1 or 2-3 flowers open 
simultaneously: flower (from base of ovary to apex of labellum) 11.5-13 
cm long. Sterile bracts 1-3; 5-6 x 1-1.5 cm; rigid, spreading slightly outwards 
from a loosely imbricating base; lanceolate, concave, apex acute, with a 
stiff, sharp point; outer surface glabrous: white. Floral bracts like sterile 
bracts, c. 5 x 1 cm. Bracteole 4.5-5 x 1.5-2 cm (when flattened); unequally 

2- lobed; apex of lobe rounded with c. 3 cm slit on opposite side; glabrous; 
red. Calyx 6.5-7 x 1.5 cm (when flattened); 3 lobed, each lobe with an 
apical tuft of long, brown hairs: tube with c. 2.5 cm slit on opposite side; 
glabrous, tawny hairs at apex only; pink, red at apex. Corolla tube 4.5-6 cm 
long; inner surface densely covered with long hairs at entrance into tube, 
rest of tube glabrous: lobes (petals) c. 1-2 cm longer than calyx, 3-4 cm 
long; apices rounded, not hooded, red; the dorsal lobe hiding the anther 
and stigma, c. 1.5 cm wide, lateral lobes narrower, c. 1 cm. Labellum held 
horizontal; blade elongate, c. 5 cm long; c. 1.5 cm wide; blade yellow with 
red margins, basal lobes yellow tinged red; margin of blade crisped and 
crinkled, apex rounded, entire or slightly retuse. Stamina! tube c. 1.5 cm 
long above the base of the corolla lobes; inner surface of the base of the 
tube densely covered with long hairs, (not warty). Filament short, c. 0.5 x 
0.6 cm wide, back glabrous. Anther c. 1 x 0.5 cm; red, hairy on pollen sacs. 
Epigynous glands 0.6-0.7 cm by 0.2-0.3 cm: comprising 2 fleshy, compressed, 
blade-like structures, not encircling base of style, each blade not further 
lobed, apex glabrous, shortly-pointed, surface not warty. Ovary c. 0.5 x 0.5 
cm, with white, short hairs; style pale-pink, hairy; stigma white, 0.3 cm 
across. Infructescence spherical; small, c. 4 x 4 cm in diameter; not 



228 



Gard. Bull. Singapore 53 (2001) 



surmounted by withered flowers; bracts completely disintegrated at fruit 
maturity; 1 — 3 fruits per head. Fruits spherical; c. 2-4 x ?>-A cm wide at the 
middle; densely covered with appressed white, short hairs; apex with scar 
of calyx only, hardly 0.1 cm long; pericarp smooth, not ridged longitudinally; 
pericarp c. 0.1 cm thick; white. Seeds irregularly angular, truncate, c. 0.6 x 
0.4 cm wide at the top. 

Distribution: Endemic to Peninsular Malaysia. Although there are few 
collections, where this species occurs, it is common. Weber (1995) mentions 
that it has been found in Selangor, Pahang, Terengganu and Johor. It has 
also been found in Perak (Khaw, 2000). 

Notes: This species is easily recognized in being few-flowered, usually with 
only 1-3 flowers. In flower coloration, it is similar to Etlingera punicea in 
that the lip has a yellow centre but it is a distinct species as the leafy 
shoots, inflorescence (Fig. IE, F), flowers (Fig. 21, J, M, N) and 
infructescence (Fig. 4, E-G, H) of E. pauciflora and E. punicea are entirely 
different. 

Specimens seen: JOHOR: Kota Tinggi. Holttum s.n. (SING); Sg. Segun, G. Panti. Corner 
SFN 30890 (2 sheets) (SING). TERENGGANU: Ulu Kajang, Kemaman. Corner SFN 
30430 (SING). 

8. Etlingera subterranea (Holttum) R.M. Sm. Notes Roy. Bot Gard. Edinb. 
43 (1986) 250. Syn.: Achasma subterraneum Holttum, Gard. Bull. Sing. 13 
(1950) 187. Type: Pahang, Cameron Highlands, 5000 ft., Holttum, SFN 
31212 (SING). 

Distribution: This species is endemic to Peninsular Malaysia. 

Notes: I have not been able to locate this species in the field. It has only 
been collected twice, on both occasions from Cameron Highlands, Pahang. 
According to Holttum (1950): "This is like A. sphaerocephalum in flowers, 
but like A. pauciflorum in inflorescence. In leaf-characters it is rather 
between A. sphaerocephalum and A. macrocheilos, having a petiole of 
moderate length. ... The fruits agree exactly with the fruits of A. 
megalocheilos as described in Java by Valeton (Ic. Bog. 2: t. 199), but are 
only 2 or 3 instead of 12 or more". 

Other specimen seen: PAHANG: Cameron Highlands, 4.600 feet, Henderson SFN 23562 
(SING). 



Etlingera (Zingiberaceae) in Peninsular Malaysia 



229 



9. Etlingera punicea (Roxb.) R.M. Sm. Notes Roy. Bot. Gard. Edinb. 43 
(1986) 250. Syn.: Alpinia punicea Roxb.. Fl. Indica 1 (1820) 73; Achasma 
macrocheilos Griff.. Notul. Plant. Asiat. 3 (1851) 429, Ic. Plant. Asiat. 
(1851) t. 357; Holttum, Gard. Bull. Sing. 13 (1950) 188; Amomum 
macrocheilos Baker. Fl. Brit. India 6 (1892) 235; Hornstedtia macrocheilos 
Ridl. in J. Str. Br. Roy. Asiat. Soc. 32 (1899) 147; Fl. Malay Penins. 4 
(1924) 271; Hornstedtia metriocheilos (Griff.) Ridl.. in J. Str. Br. Roy. 
Asiat. Soc. 32 (1899) 147; Fl. Malay Penins. 4 (1924) 271 (not of Griffith). 
Type: Malacca. Griffith s.n., s.v. (specimen lost?). 

Figure IF, 2M & N, 31, 4H 

Rhizomes to c. 12 cm below ground; slender, c. 1.5 cm diam.; overlapping 
scale-leaves, green tinged dull red or pink. Leafy shoots 2-5 m tall; close 
together or widely spaced. 1 1^40 cm apart: sheaths conspicuously ribbed 
with cross bars covered by tufts of short, white hairs (clearer in young 
sheaths), sometimes smooth, green or yellowish with purple blotches, 
crushed sheaths without distinct scent; ligule 0.5-1 cm. apex broadly 
rounded, entire, margin and surface densely covered by short, brown hairs; 
petiole c. 0.5 cm or subsessile. surface reticulate. Laminas c. 10 pairs; 
(topmost) c. 58 x 9 cm, (middle) 62-91 x 14-17 cm. (basal) 18-22 x 7 cm: 
upper surface smooth or with slightly raised lateral veins: entirely green 
(sometimes with brown bars on lower surface of young leaves): lower 
surface glabrous; base cuneate. sometimes with auricles. Peduncle 5-10 
cm; horizontal, then erect, entirely subterranean at base of leafy-shoot or 
some distance away; sheaths c. 13. shortest at base 0.5-5 cm long, apex 
acute, with a stiff spine 0.4 cm long, overlapping, pale green. Inflorescence 
entirely subterranean except for apices of sterile bracts and flowers. Fertile 
apical part of inflorescence with well developed sterile bracts tightly 
imbricating forming a narrow sub-cylindric structure. 10-13 x c. 4 cm wide, 
in side view hiding c. 2/3 or more of the inflorescence head proper: sterile 
bracts persisting to fruit maturity. Receptacle of inflorescence c. 0.5 cm 
long: apex inflated and rounded; bearing c. 30 flowers; 7-10 flowers open 
simultaneously: flower (from base of ovary to apex of labellum) 10.5-12.5 
cm long. Sterile bracts c. 5; 5.5-6 x 2-2.5 cm; rigid, erect, lanceolate, broadest 
at middle, strongly concave; apex acute, with a stiff, sharp point; outer 
surface sparsely covered with white hairs, more densely hairy at base; 
white, pink towards apex. Outer floral bracts resembling the sterile bracts; 
c. 6x2 cm; inner floral brads smaller c. 5 x 1 cm. Bracteole c. 4 x 1.5 cm 
(when flattened): 2-lobed, apex of lobe rounded with 1.5 cm slit on opposite 
side; surface with light brown hairs; pink. Calyx c. 7 x 1.5 cm (when 
flattened); 3-lobed. each lobe with a hairy, subapical tooth c. 0.1 cm; tube 
with c. 3 cm slit on opposite side; glabrous, with tawny hairs only at apex. 



230 



Gard. Bull Singapore 53 (2001) 



Corolla-tube c. 6 cm; inner surface with hairs mostly at upper 1/3 of tube; 
lobes (petals) c. 1-2 cm longer than calyx, c. 3 cm long, apices rounded, not 
hooded, red; dorsal lobe hiding the anther and stigma, c. 0.8 cm wide, 
lateral lobes narrower, c. 0.5 cm. Labellum held horizontal, blade elongate, 
5.5-6 cm long, c. 1.5 cm wide; blade and basal lobes yellow with a narrow 
scarlet margin broadening to the spatulate apex, scarlet margins of the haft 
infolded so that the 'margin' appears to be yellow; margin of blade crisped 
and crinkled; apex rounded, retuse or deeply bilobed. Stamina! tube 0.8-1 
cm long above the base of the corolla-lobes; inner surface of tube glabrous 
or sparsely covered with long hairs, (not warty). Filament short, c. 0.3 x 0.3 
cm, adaxially glabrous. Anther c. 1 x 0.5 cm; red, hairy on pollen sacs. 
Epigynous glands c. 0.5 x 0.2-0.3 cm; comprising 2 fleshy compressed blade- 
like structures, not encircling base of style, each blade not further lobed, 
apex glabrous, shortly-pointed, surface not warty. Ovary c. 0.5 x 0.4 cm; 
covered with appressed, white, short hairs; style white, hairy; stigma pink, 
0.3 cm across. Infructescence globular; large c. 6 x 9 cm in diameter; not 
surmounted by withered flowers; sterile bracts at base of fruit head persisting 
to fruit maturity; floral bracts persistent with fruits at their axils; 5 — 15 
fruits per head. Fruits obovoid; 3-3.5 x 3^4.5 cm wide at the top; densely 
covered with appressed short hairs; apex with remnant of calyx base only, 
less than 0.5 cm long; pericarp ridged longitudinally, each ridge with many 
irregular, coarse, spine-like processes; pericarp c. 0.2 cm thick (excluding 
ridges), pinkish-red. Seeds irregularly angular, truncate; c. 0.4 x 0.3 cm 
wide at the top. 

Distribution: A common species found in lowland forests throughout the 
Peninsula. It also occurs in Thailand, Borneo, Java and Sumatra (Smith, 

1986). 

Notes: The conspicuous character of this species is the colour of the lip. 
yellow in the middle, the rest scarlet. There is variation in the details of the 
lip, especially the tip of the mid-lobe. 

Specimens seen: PERAK: Tanjong Malim, Burkill & Haniff 13476 (SING). SELANGOR: 
Genting Bidai, Ridley 7812 (SING); Genting Simpah, Hume 9722 (SING); Klang, Teluk 
Reserve, Burkill S FN 5997, Bukit Rajah Forest, Burkill 15512 (SING); Ulu Gombak. Yong 
FRI 99256 (KEP). NEGRI SEMBILAN: G. Tampin. Burkill 3179 (SING); Seremban, G. 
Telapak Buruk, Saw FRI 40052 (KEP). JOHOR: B. Tinjau Laut. Corner SFN 37061 (2 
sheets) (SING): Sg. Pelepah, G. Panti, Md. Nur 20012 (SING); Kuala Tebing Tinggi. Ridley 
s.n. (SING); Batu Pahat. Ridley s.n. (SING); Sg. Kayu Ara, Mawai-Jemaluang Rd.. Corner 
SFN 31476 (SING); G. Pulai. Ridley s.n. (SING); G. Panti, Ridley s.n. (SING); 12 Mile 
Mawai-Jemaluang Road, Corner SFN 29993 (SING); Sg. Tebrau. Ridley s.n. (SING). 
TERENGGANU: Sg. Kemaman, Kampong Ayer Puteh, Corner SFN 307/7 (KEP. SING). 



Etlingera < Zingiberaceae) in Peninsular Malaysia 



231 



10. Etlingera metriocheilos (Griff.) R.M. Sm. Notes Roy. Bot Gard. Edinb. 
43 (1986) 247. Syn.: Achasma metriocheilos Griff.. Notul. Plant. Asiat. 3 
(1851) 427. Ic. Plant Asiat. (1851) t. 356: Amomum sphaerocephalum Bak. 
in Hook. /.. Fl. Brit. India 6 (1892) 234: Amomum metriocheilos Bak. in 
Hook. /.. Fl. Brit. India 6 (1892) 234: Hornstedtia albomarginata RidL in 
J. Str. Br. Roy. Asiat. Soc. 32 (1899) 145: Fl. Malay Penins. 4 (1924) 269: 
Hornstedtia sphaerocephala (Baker) K. Schum.. Pflanzenr. Zingib. (1904) 
192: Achasma sphaerocephalum (Bak.) Holtt.. Gard. Bull. Sing. 13 (1950) 
189. Type: Malacca. Griffith 5758, s.v. (K). 

Figure 1G, 2K & L 3J, 41 

Rhizomes just below ground or creeping at soil surface: stout, 2-4.5 cm 
diam.: scale leaves overlapping, conspicuously pink, bright red or dull 
purple-red. Leafy shoots 2-6 m tall: close together or widely spaced. 10-80 
cm apart: sheaths dull red or green, conspicuously ribbed longitudinally 
with cross-bars covered with tufts of short, white hairs or with sheaths 
bright red, surface slightly reticulate: crushed sheaths without distinct scent; 
ligule c. 1 cm long, apex broadly rounded, entire, margin and surface with 
short, brown hairs (densely, silky-hairy in var. major): petiole 0.5-1.5 cm 
(c. 3.5 cm in var. petiolata). surface reticulate. Laminas c. 15 pairs: (topmost) 
22-55 x 3-^.5 cm, (middle) 57-80 x 9.5-14.5 cm. (basal) 9-25 x 5.5-8.5 cm; 
smooth (prominently ribbed in var. petiolata): entirely dark green above, 
when young dark purple-red beneath (or with brown-red bars on upper 
surface in var. rub ro striata): lower surface glabrous (with rather rough, 
short, brown hairs in var. major): base variable on the same leafy shoot, 
rounded or cuneate. Peduncle 4-10 cm: horizontal, then erect, entirely 
subterranean at base of leafy shoot or more usually some distance away: 
sheaths c. 7. shortest at base 1-4 cm long, apex acute with subapical spine 
to 0.2 cm. overlapping, red. Inflorescence entirely subterranean except for 
apices of sterile bracts and flowers. Fertile apical part of inflorescence with 
well developed sterile bracts tightly imbricating forming a narrow sub- 
cylindric structure 4-6 x c. 5 cm. hiding c. 1/3 of the inflorescence head 
proper in side view: sterile bracts persisting to fruit maturity. Receptacle of 
inflorescence c. 2 cm long: apex inflated and conical: bearing 25-60 flowers: 
5-6 flowers open simultaneously: flower (from base of ovary to apex of 
labellum) 8-10.5 cm long. Sterile bracts c. 8; 3-4.5 x 2-3 cm: rigid, erect, 
oblong-elliptic, broadest below middle, strongly concave; apex acute, with 
a stiff spine 0.2 cm long: outer surface densely covered with appressed, 
brown hairs, hairs golden brown at base of bract; entirely red, or red at 
base and deep purple-red at apex. Outer floral bracts resembling the sterile 
bracts, c. 4.5 x 1.5 cm; inner floral bracts smaller, c. 4 x 1 cm. Bracteole 2.5- 
3 x c. 1.5 cm (when flattened); unequally 2-lobed. apex of lobe acute with 



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Card. Bull. Singapore 53 12001) 



c. 1 cm slit on opposite side; surface with brown hairs, red. Calyx 5-6.5 x 
1.5 cm (when flattened); 3-lobed, each lobe with a glabrous, subapical 
tooth c. 0.2 cm long; tube with 2.5-3 cm slit on opposite side; glabrous, 
only the apex with tawny hairs; red. Corolla tube 2.5-3.5 cm long; inner 
surface of tube at the entrance sparsely hairy, rest of tube glabrous; lobes 
(petals) variable: equal to, shorter or longer than calyx, 2-2.5 cm long, 
apices rounded, not hooded, red; dorsal lobe not hiding the anther and 
stigma, c. 0.5 cm wide, lateral lobes narrower, c. 0.4 cm. Labellum held 
horizontal, blade elongate, 3.5-5 cm long, c. 1.5-1.8 cm wide; blade and 
basal lobes crimson, margins of lobes white; margin of blade plain or slightly 
crisped; apex retuse or deeply bilobed. Staminal tube 1.5-2 cm long above 
the base of the corolla-lobes, inner surface of tube glabrous or sparsely 
covered with long hairs, (not warty). Filament short, c. 0.6 x 0.3 cm wide, 
adaxially glabrous. Anther c. 1 x 0.5 cm; crimson, hairy on pollen sacs. 
Epigynous glands 0.3-0.4 x 0.3 cm; comprising 2 rounded mounds, 
surrounding base of style, each mound minutely lobed, apices hairy, surface 
not warty. Ovary c. 0.5 x 0.4 cm; densely covered with appressed, white, 
short hairs; style pink, hairy; stigma dark purple, almost black, 0.4 cm 
across. Infructescence globular; large, c. 7 x 9 cm diam.; not surmounted by 
withered flowers; sterile bracts at base of fruit head and floral bracts 
persisting to fruit maturity; 25 — 45 fruits per head. Fruits obovoid; 3-4 x c. 
2.5 cm wide at the top; densely covered with appressed, short hairs; apex 
with remnant of the base of the calyx only, less than 0.5 cm long; pericarp 
ridged longitudinally, each ridge with a shoulder-like apical mound, covered 
in prominent warts; pericarp c. 0.2 cm thick (excluding ridges); dark purple. 
Seeds irregularly angular, truncate, c. 0.4 x 0.3 cm wide at the top. 

Distribution: Found in many parts of the Peninsula, in lowland forests and 
at moderate elevations on the mountains. It is also found in Borneo (Smith, 

1986). 

Notes: This species is variable and Holttum (1950) described four varieties, 
ranging from the few-flowered, small-sized var. rubrostriata, to the robust 
var. petiolata. The distinguishing characters of this species are: (1) the 
striking red or purple lower surface of young leaves and leaf-sheaths except 
for var. rubrostriata, which has a red-barred upper surface and is green 
beneath, (2) a sub-globose or sub-cylindric inflorescence, (3) the deep red 
or crimson involucral bracts that are distinctly shorter than the calyces, 
and (4) the deep crimson lip with white (never yellow) margins. 

While noting it was possible that Achasma sphaerocephalum (Baker) 
Holttum was identical with A. metriocheilos Griff., Holttum (1950 p. 191) 
nevertheless thought it better to exclude Griffith's name. The reason he 



Etlingera (Zingiberaceae) in Peninsular Malaysia 



233 



cited was that the details given by Griffith were inadequate for him to 
decide with certainty whether the two species were conspecific. 

Smith (1986 p. 247) combined Achasma metriocheilos Griff, with 
Amomum sphaerocephalum Bak. as Etlingera metriocheilos (Griff.) R.M. 
Sm.. However, she included in the combination. Hornstedtia metriocheilos 
(Griff.) RidL the species which Ridley (1899 p. 147) described as having a 
crimson lip with a yellow centre. However, this is a different species and. 
as Holttum had noted, it is in fact Etlingera punicea. 

Types seen: Syntypes of Hornstedtia albomarginata RidL. J. Str. Br. Roy. 
Asiat. Soc. 32 (1899) 145. — Penang. road to Penara Bukit. Ridley s.n. 
(SING): Selangor. Petaling. Ridley s.n. (SING). 

Achasma sphaerocephalum (Bak.) var. petiolatum Holttum. in Gard. Bull. 
Sing. 13 (1950) 191. — Johor, Ulu Segun. G. Panti. Corner SFN 30745 
(SING). 

var. rubrostriatum Holttum. in Gard. 
Bull. Sing. 13 (1950) 190. — Selangor. K. Kubu side of the Gap. Corner 
SFN 30776 (SING). 

var. grandiflorum Holttum. in Gard. 
Bull. Sing. 13 (1950) 191. — Terengganu. Kemaman. Bukit Kajang. Corner 
SFN 30234 (SING). 

var. majus Holttum. in Gard. Bull. Sing. 
13 (1950) 190. — Terengganu. Kemaman. Bukit Kajang. Corner SFN 30205 
(SING). 

Other specimens seen: PERAK: Maxwell's Hill. Coll. ? s.n. (SING): Tea Gardens. Curtis s.n. 
(SING): Kuala Kangsar. Kg. Hitam Jong. Sg. Plus F.R.. Hashim Pendek KLU 119 (KEP). 
SELANGOR: Petafing. Ridley s.n. (SING): UPM campus. Anthonysamy SA 199 (KEP). 

11. Etlingera triorgyalis (Baker) R.M. Sm. Notes Roy. Bot. Gard. Edinb. 
43 (1986) 250. Amomum triorgyale Baker, in Hook. f.. Fl. Brit. India 6 
(1892) 237: Hornstedtia triorgyale (Baker) Ridl.. in J. Str. Br. Roy. Asiat. 
Soc. 32 (1899) 144: Mat. Fl. Malay. Penins. 1 (1907) 38: Fl. Malay Penins. 4 
(1924) 269; Achasma triorgyale (Baker) Holttum. Gard. Bull. Sing. 13 (1950) 
186. Type: Perak. Larut. King's Collector 2105, s.v. (K). 

Figure ID, 2G & H, 3G, 4D 

Rhizomes just below ground: stout. 2.5-4.5 cm diam.: scale leaves 
overlapping, green tinged dull red or pink. Leafy shoots 5-6 m tall: close 
together or widely spaced to c. 1 m apart: sheaths conspicuously ribbed 
longitudinally with cross-bars covered by tufts of short, white hairs 
(especially in young sheaths), when crushed with a strong scent suggestive 



234 



Card. Bull. Singapore 53 (2001) 



of pepper; ligule 1-2.5 cm; apex rounded, entire; margin and surface with 
short, brown hairs; petiole 1.5-3 cm, surface reticulate. Laminas c. 15 pairs; 
(topmost) c. 43 x 3.5 cm, (middle) 80-100 x 6-21.5 cm, (basal) c. 25 x 7 cm; 
smooth or with slightly raised lateral veins, when young entirely green on 
both surfaces; lower surface softly hairy with short hairs; margin broadly 
crisped, red with brown hairs; base broadly cuneate, unequal, slightly 
decurrent. Peduncle short, 5-15 cm; horizontal, then erect, entirely 
subterranean at base of leafy shoot; sheaths c. 14, shortest at base 4-7.5 cm 
long, apex acute with subapical spine 0.2-1 cm long, white or pale green 
tinged deep pink, overlapping. Inflorescence subterranean except for distal 
parts of sterile bracts and flowers. Fertile apical part of inflorescence with 
well developed sterile bracts tightly imbricating, forming a narrow sub- 
cylindric structure 10-14 x c. 6 cm wide, in side view hiding c. 2/3 or more 
of the inflorescence-head proper, the sterile bracts not persisting to fruit 
maturity. Receptacle of inflorescence c. 1 cm, apex inflated and rounded; 
bearing 40-50 flowers; 8-17 flowers open simultaneously; flower (from 
base of ovary to apex of labellum) c. 13.5 cm long. Sterile bracts c. 7; 9-11 x 
3-6 cm; rigid, erect, oblong-obovate, recurved at apex, concave, apex 
broadly rounded, retuse, with subapical spine c. 0.1 cm; outer surface with 
white hairs, hairs densely appressed at base; white or pale green, deep 
pink towards apex. Outer floral bracts resembling the sterile bracts, c. 10.5 
x 2.5 cm; inner floral bracts smaller, c. 8.5 x 1 cm. Bracteole c. 8 x 2 cm 
(when flattened); 2-lobed, apex of lobe rounded with c. 2 cm slit on opposite 
side; surface with light-brown hairs; white with red apex. Calyx 7.5-9 x c. 2 
cm (when flattened); 3-lobed, each lobe with a hairy, subapical tooth c. 0.1 
cm long; tube with c. 4.5 cm slit on opposite side; glabrous, with tawny- 
hairs at apex only; white, pink-red at apex. Corolla tube 5.8-6.5 cm long, 
inner surface densely covered with long hairs at entrance of tube, rest of 
tube glabrous; lobes (petals) equal to calyx, c. 2.8 cm long, apices rounded, 
not hooded, red; dorsal lobe not hiding the anther and stigma, c. 1 cm 
wide, lateral lobes narrower, c. 0.7 cm. Labellum held horizontal, blade 
elongate, c. 5 cm long, c. 2.5 cm wide; blade and basal lobes entirely bright 
pink-red; margin of blade crisped and crinkled, apex broadly rounded and 
retuse. Staminal tube c. 1.2 cm long above the base of the corolla lobes; 
inner surface of tube densely covered with long hairs below anther only, 
rest of surface densely covered by warts. Filament short, 0.4 x 0.3 cm, 
adaxially hairy. Anther c.l x 0.5 cm, red. Epigynous glands c.l x 0.2-0.3 cm; 
comprising 2 fleshy compressed blade-like structures, not encircling base 
of style, each blade not further lobed, apex glabrous, shortly-pointed, surface 
not warty. Ovary c. 1 x 0.5 cm, densely covered with appressed, white short 
hairs; style pale pink, hairy; stigma pale pink, c. 0.5 cm across. Infructescence 
globular, large, c. 6 x 10 cm diam.; not surmounted by withered flowers; 



Etlingera (Zingiberaceae) in Peninsular Malaysia 



235 



bracts completely disintegrated at fruit maturity; 20 — 35 fruits per head. 
Fruits obovoid; c. 3 x 2.5-3 cm wide at the top: densely covered with 
appressed, short hairs; apex with remnant of calyx base only, less than c. 
0.5 cm long; pericarp ridged longitudinally, each ridge with many irregular, 
fine spine-like processes; pericarp c. 0.2 cm thick (excluding ridges), dark 
red or pink. Seeds irregularly angular, truncate c. 0.4 x 0.3 cm wide at the 
top. 

Distribution: In Peninsular Malaysia, recorded from Perak, Selangor and 
Pahang. It also occurs in Borneo and Sumatra (Smith. 1986). 

Notes: The very broad bracts, recurved at the deep rose apices, together 
with the cherry red or deep rose lips are striking characters of this species. 

Specimens seen: PERAK: Ipoh, foot of limestone rocks, Curtis SFN 3317 (SING). 
SELANGOR: Genting Peras. Ridley 7806 (SING). PAHANG: Ulu Sg. Sat, Mohd. Shah & 
Mohd. Noor MS 1880 (KEP. SING): Temerloh. Sg. Nering. Henderson 10583 (SING). 

12. Etlingera littoralis (Konig) Giseke. Syn.: Hornstedtia megalochilus Ridl.. 
in J. Str. Br. Roy. Asiat. Soc. 32 (1899) 146; Mat. Fl. Malay. Penins. 1 
(1907) 38; Fl. Malay Penins. 4 (1924) 270: Amomum megalocheilos Baker, 
Fl. Brit. India 6 (1892) 236; Achasma megalocheilos Griff., Notul. Plant. 
Asiat. 3 (1851) 426, Ic. Plant. Asiat. (1851) t. 355; Valeton., Ic. Bog. 2 
(1903) t. 188, 199. 

Figure 1H, 20 & P, 3K 

Rhizomes at surface or below ground to c. 10 cm deep; thick, 3-3.5 cm 
diam.: scale-leaves overlapping, green tinged dull red. Leafy shoots 3-6 m 
tall; close together or widely spaced, 12^48 cm apart; sheaths conspicuously 
ribbed longitudinally, with cross-bars covered by tufts of short, white hairs 
(clearer in young sheaths); crushed sheaths without distinct scent; ligule 
1.5-2 cm long; apex broadly rounded, entire; margin with short brown 
hairs, surface with purple blotches, glabrous; petiole 1.5^1.5 cm; surface 
reticulate. Laminas c. 13 pairs; (topmost) c. 69 x 8 cm, (middle) c. 101 x 18 
cm, (basal) c. 27 x 7 cm; smooth; entirely green, in young leaves sometimes 
flushed pink beneath; mostly glabrous, sometimes with short hairs; base 
variable on the same leafy shoot, cuneate, rounded or truncate, always 
unequal. Peduncle 4-10 cm; horizontal, then erect, entirely subterranean, 
at base of leafy shoot or some distance away: sheaths c. 11, shortest at base 
from 1.5-6 cm. apex acute with a stiff subapical spine 0.1 cm, overlapping, 
pale green or white tinged pink. Inflorescence subterranean except for 
apices of sterile bracts and flowers. Fertile apical part of inflorescence with 



236 



Card. Bull. Singapore 53 (2001) 



well developed sterile bracts tightly imbricating forming a narrow sub- 
cylindric structure c. 12 x 5 cm, in side view hiding c. 2/3 or more of the 
inflorescence-head proper. Receptacle of inflorescence c. 0.5 cm; apex 
inflated and rounded; bearing 30-35 flowers; 5-9 flowers open 
simultaneously; flower (from base of ovary to apex of labellum) 13-16 cm 
long. Sterile bracts 5 or 6; 6-8.5 x 2-5 cm; rigid, erect, oblong-obovate, 
broadest at middle, concave; apex acute, with a stiff spine 0.1 cm long; 
outer surface glabrous, densely covered with appressed. white hairs at base 
only; white, pink towards apex. Outer floral bracts resembling the sterile 
bracts; c. 8.5 x 1.5-2.5 cm; inner floral bracts smaller, c. 8 x 0.5 cm. Bracteole 
7-7.5 x c. 1.5 cm wide (when flattened); 2-lobed, apex of lobe acute with 3 
cm slit on opposite side; surface with brown hairs; pale pink. Calyx 7-9 x c. 
1.5 cm (when flattened); 3-lobed; subapical tooth inconspicuous; tube with 
c. 3 cm slit on opposite side; glabrous, with tawny hairs at apex only; pink. 
Corolla tube 6-6.8 cm long, inner surface densely covered with long hairs 
at entrance oftube, rest of tube sparsely hairy; lobes (petals) equal to calyx, 
c. 2.5 cm long, apices rounded, not hooded, pink; dorsal lobe not hiding 
the anther and stigma, c. 1 cm wide, lateral lobes narrower, c. 0.5 cm. 
Labellum held horizontal, blade elongate, c. 7 cm long, c. 2 cm wide; blade 
and basal lobes entirely red or with a yellow margin sometimes not reaching 
the broad apex; margin of blade plain or crisped and crinkled; apex rounded, 
entire or slightly retuse. Staminal tube c. 1.5 cm long above the base of the 
corolla-lobes; inner surface of tube densely covered with long hairs, (not 
warty). Filament short, c. 0.5 x 0.4 cm wide, adaxially hairy. Anther c. 1 x 
0.5 cm; red, hairy on pollen sacs. Epigynous glands c. 1 x 0.2-0.3 cm; 
comprising 2 fleshy, compressed, blade-like structures, not encircling base 
of style, each blade not further lobed, apex glabrous, shortly pointed, surface 
not warty. Ovary 0.5-0.6 x 0.5 cm; densely covered with appressed, yellow, 
short hairs; style pink, hairy: stigma red. 0.3 cm across. Fruit not known. 

Distribution: Common in lowland forests throughout the Peninsula. It is 
also found in China, Thailand. Borneo, Java and Sumatra (Smith, 1986). 

Notes: There is variation in lip colour in this species. Most commonly 
encountered are plants with scarlet lips with a yellow margin. However, 
plants growing nearby may have scarlet lips edged orange-red or a third 
type with much narrower lips are scarlet with a thin yellow margin. 

The description of the infructescence by Holttum (1950) was based 
on fruits described from Java by Valeton. I have attempted in the field to 
find the infructescence of this common species but without success. The 
inflorescences in a large patch growing in the Temenggor F.R., Perak, 



Etlingera (Zingiberaceae) in Peninsular Malaysia 



237 



were tagged and the site revisited two months later to look for the fruits 
but none was found. The fact that it has never been found in fruit in 
Peninsular Malaysia warrants further investigation. 

Specimens seen: PERLIS: Wang Kelian. Wan Fadhilah HI 919 (KLU); Mata Ayer F.R.. 
Saw, Kamarudin, Jamaludin & Baya FRI s.n. (KEP). PERAK: Tambun limestone cliff, 
Burkill SFN 6294 (SING); Upper Perak, Wray 3444 (SING); SELANGOR: Gua Batu. 
Ridley s.n. (SING); Kuala Lumpur. Ridley s.n. (SING). MALACCA: Bukit Sadanen, Ridley 
s.n. (SING). JOHOR: Castleroad. Ridley s.n. (SING); Sg. Segun, G. Panti. Corner SFN 
30891, 30892 (SING); Batu Pahat, Ridley s.n. (SING); Kota Tinggi. Ridley s.n. (SING). 
KELANTAN: Sg. Lebir. Ben. Stone & Mahmud Sidek 12507 (KEP). PAHANG: Jerantut, 
Corner SFN 30768 (SING); Temerloh. Titi Bungor. Henderson 10674 (SING): Telom, Ridley 
13832 (SING): Tanjong Gajah, Ridley s.n. (SING): Sg. Pertang. Bentong, Burkill & Haniff 
SFN 16515 (SING): Pekan, Ridley 1632 (SING); Tahan Woods. Ridley s.n. (SING). 
TERENGGANU: Kemaman. Sg. Nipa. Corner SFN 30581 (SING), Bukit Kajang. Corner 
SFN 30378 (SING). 

Acknowledgements 

I thank Dr Saw Leng Guan. (KEP) for his encouragement, advice and 
providing photographs of the type and other specimens of Phaeomeria 
venusta from Kew and to the Curators of KEP, KLU and SING for 
permission to use their herbaria. I thank Dr J. Dransfield (Royal Botanic 
Gardens. Kew) for the Latin diagnosis. I also acknowledge with thanks Dr 
Saw Leng Guan. Dr Ruth Kiew and unnamed reviewers for advice on this 
paper. Thanks are due to the Malaysian Nature Society for help during the 
Malaysian Heritage and Scientific Expedition: Belum phase 1 (1993/1994) 
and phase 2 (1998). Advice from Dr E. Soepadmo (FRIM) during the 
Belum Expeditions is also gratefully acknowledged. Help and 
companionship in the field from Ms Rosema Abdullah and Mr Abdullah 
Piee are very much appreciated. 

References 

Burtt. B.L. and R.M. Smith. 1986. Etlingera: The inclusive name for 
Achasma, Geanthus and Nicolaia (Zingiberaceae). Notes Royal Botanic 
Garden Edinburgh. 43: 235-241. 

Holttum. R.E. 1950. The Zingiberaceae of the Malay Peninsula. Gardens' 
Bulletin, Singapore. 13: 1-249. 

Ibrahim. H. 1986. Notes on gingers of Pulau Tioman. Nature Malaysiana. 
11(4): 10-13. 



238 



Card. Bull Singapore 53 (2001) 



Khaw, S.H. 2000. A checklist of the Gingers (Zingiberaceae) of the Belum 
and Temenggor Forest Reserves in Perak, Peninsular Malaysia. Malayan 
Nature Journal. 54: 227-232. 

Larsen, K. 2000. Selection of Etlingera corneri J. Mood & H. Ibrahim 
(Zingiberaceae) as a future correct name. Nordic Journal of Botany. 20: 
475-476. 

Larsen, K., H. Ibrahim, S.H. Khaw and L.G. Saw. 1999. Gingers of 
Peninsular Malaysia and Singapore. Natural History Publications 
(Borneo), Kota Kinabalu, Sabah. Malaysia. 135 pp. 

Larsen, K.1970. The genus Nicolaia in Thailand. Natural History Bulletin 
Siam Society. 23: 574-575. 

Lim, C.K. 2000. Taxonomic notes on Etlingera Giseke (Zingiberaceae) in 
Peninsular Malaysia: the "Nicolaia" taxa. Folia malaysiana. 1: 1-12. 

Mood, J. & H. Ibrahim. 2000. A new species of Etlingera (Zingiberaceae) 
from Peninsular Malaysia and southern Thailand. Nordic Journal of 
Botany. 20: 279-283. 

Ridley, H.N. 1899. The Scitamineae of the Malay Peninsula. Journal Straits 
Branch Royal Asiatic Society. 32: 85-184. 

Ridley, H.N. 1924. Zingiberaceae. Flora of the Malay Peninsula. 4: 233- 
285. Reeve & Co., London. 

Smith, R.M. 1986. New combinations in Etlingera Giseke (Zingiberaceae). 
Notes Royal Botanic Garden Edinburgh. 43: 243-254. 

Weber, A. 1995. Etlingera gingers (Zingiberaceae) of Peninsular Malaysia. 
Nature Malaysiana. 20(3): 80-89. 

Postscript: While the account above was in press, the note by Latiff (2001) 
appeared discussing the question of priority of E. terengannuensis C.K. 
Lim and E. corneri J. Mood & H. Ibrahim. As noted above, not all the 
specimens cited by Mood & Ibrahim (2000) belong to a single taxon and 
Lim confirms that Corner SFN 32778 and "the Curtis specimen of 1890" 
belong to E. fulgens based on the purple undersides of their leaves. 
However, as Latiff points out, this does not affect the validity of the name 
E. corneri, which is based on the type so the fact that the specimens cited 
included more than one taxon is not relevant to the problem of the priority 
of names. 



Ellingera (Zingiberaceae) in Peninsular Malaysia 



239 



Detailed examination of the species in question based on living 
material has allowed accurate description of the species (see main text) 
based only on specimens of E. corneri. 

Reference: Latiff. A. 2001. Notice of priority of E. terengganuensis C.K. 
Lim. Folia malaysiana. 2(2): 75 — 68. 



Gardens Bulletin Singapore 53 (2001) 241-286. 



The Limestone Begonias of Sabah. Borneo - Flagship 
Species for Conservation 

RUTH KIEW 
Singapore Botanic Gardens. 1 Cluny Road. Singapore 259569 

Abstract 

The 18 Begonia ( Begoniaceae ) species now known from limestone and associated substrates 
in Sabah. Malaysia, are listed and a key provided for their identification. Twelve are described 
as new species: Begonia anthonyi Kiew. B. baturongensis Kiew. B. berhamanii Kiew. B. 
diwolii Kiew. B. heliosrrophe Kiew. B. keeana Kiew. B. lambii Kiew. B. layang-layang 
Kiew. B. madaiensis Kiew. B. melikopia Kiew. B. punbatuensis Kiew and B. unmensis 
Kiew. All these new species belong to sect. Petermannia (Klotzsch.) A.DC. except for B. 
diwolii that belongs to sect. Diploclinium (Lindl.) A.DC- Reasons for placing B. amphioxus 
Sands in sect. Petermannia and B. gueritziana Gibbs in sect. Platycenmun (Klotzsch.) A.DC. 
are given. Begonia is the most speciose genus on limestone in Sabah. Of the 18 species, 
only one. B. gueritziana. is widespread and also found on non-limestone substrates. Of the 
others. 12 species are known from single limestone hills and 5 are from hills within the 
same area. Habitat disturbance, which increases the risk of fire, therefore poses the greatest 
threat to the conservation of these species. 

Introduction 

As flagship species for the conservation of the limestone flora of Sabah. 
begonias meet all the criteria. Most are readily recognised as relatives of 
the begonia house plants, the majority of the limestone species are attractive 
(Kiew. 2000) and therefore have not only aesthetic appeal but also have 
commercial potential in horticulture as ornamental plants, as well as 
exemplifying the need to conserve the limestone flora to ensure its continued 
existence. 

Begonias are instantly recognised by their leaves, which are usually 
very asymmetric. Although the Sabah species have relatively small flowers, 
they make up for this deficiency either in their decorative habit or the 
great variety of leaves. Some are small rosette plants, like B. gueritziana 
and B. lambii: others are tall cane-like plants, like B. keiihii Kiew with 
bright red. glossy stems: some have leaves coloured magenta underneath, 
like B. keeana: or have pink spots, like B. malachosticta Sands, or purple- 
magenta hairs, like B. lambii: others have unusually shaped leaves, such as 
the swallow begonia. B. layang-layang. with leaves curved like a swallow s 
wing, or B. amphioxus with bizarre, narrow peltate leaves pointed at both 
ends. 



242 



Gard. Bull. Singapore 53 (2001) 



These attractive or unusual-looking species hold potential as 
ornamental plants and several have been successfully grown locally in the 
Agricultural Park at Tenom under Anthony Lamb's supervision. 

That the limestone flora in Sabah is under threat is not in doubt. The 
major threat is from accidental fires that occur in the periodic drought 
periods, which are particularly severe in El Nino years. Of the 59 limestone 
hills in Sabah, only 22 fall within protected forest or a Virgin Jungle Reserve 
(Lim & Kiew, 1997). The other hills are either no longer surrounded by 
forest or else lie within commercial forest that has been, is or will be 
logged. The vegetation on these hills is particularly vulnerable to fire. 
Beaman et ai, (1985) showed that in the Great Burn of 1982/83, the 
frequency of burning was five times greater in logged-over forest than 
undisturbed forest. Indeed, the limestone vegetation that burned then, for 
example, on Bukit Dulong Lambu (Gomantong Cave), Bukit Batangan 
and Gunung Madai, has not yet recovered. 

Other hills are under threat from quarrying as a source of cement, 
such as Pulau Balambangan and Bukit Tengar (Segarong Cave), or for 
marble, as at the Borneo Marble Quarry on the Segama River, or for road 
metal, for example, Batu Pang, Supu and Temambong. Yet others suffer 
disturbance from birdnest collecting activities, such as Gunung Madai and 
Bukit Dulong Lambu, where the latter has a village built on the summit. A 
few have become tourist attractions and are suffering degradation, such as 
Batu Tulug (Batu Putih) and Batu Punggul. Even protected status does 
not necessarily ensure protection as is demonstrated by the Virgin Jungle 
Reserve surrounding Bukit Baturong that has been logged twice (illegally) 
in the last ten years. 

Conservation of the limestone flora therefore requires a strategy to 
protect its biodiversity and Begonia serves as a useful indicator as it is one 
of the most biodiverse groups of the limestone flora in Sabah being one of 
the ten most speciose families and the most speciose genus (Kiew, 1998a). 
In addition, it exhibits extremely high endemism. All its 18 species are 
endemic to Sabah. Only one species, B. gueritziana, is widespread; the rest 
are narrow endemics. Twelve species are found on single hills and five 
species on adjacent hills (Table 1). In addition, their distribution reflects 
the general regional differences of the limestone flora as a whole in Sabah 
because their distribution is mirrored by the distribution of other species 
(Kiew, 1998a). Therefore, by using the distribution of begonia species, a 
pattern emerges of those hills that are of greatest importance in order to 
protect the maximum biodiversity of the limestone flora. (The only 
limestone localities without begonia species are the northern islands of 
Balembangan and Banggi). Begonia is therefore an obvious choice as a 
conservation icon to illustrate the need for conserving a network of hills in 



Limestone Begonias ofSabah. Borneo 



243 



order to include maximum biodiversity 

Table 1. Distribution of Begonia species on limestone in Sabah. 



District 



Locality 



Begonia Species 



(a) species found on single hills 



Pensiangan 
Pensiangan 
Pensiangan 
Pensiangan 
Kinabatangan 

Kinabatangan 
Lahad Datu 
Lahad Datu 



Batu Punggul/Batu Tinahas 
Sapulut 
Batu Urun 
Pun Batu 

Bukit Dulung Lambu 

Melikop 
Gunung Madai 
Bukit Baturong 



Semporna Batu Tengar 

(b) species found on several hills in the same area 

Pensiangan Batu Punggul/Tinahas. Sapulut 

Kinabatangan Kinabatangan Valley (3 hills) 

Kinabatangan Kinabatangan Valley (2 hills) 

Lahad Datu Segama River (5 hills) 



amphioxus, anthonyi 

layang-layang 

urunensis 

punbatuensis 

gomantongensis, 

malachosticta 

melikopia 

madaiensis 

baturongensis, 

berhamanii 

keithii 



lambii 
postarii 
heliostrophe 
diwolii, keeana 



(c) widespread on limestone in Sabah 



gueritziana 



Two hills are outstanding for their variety of begonia species. Both 
Bukit Dulong Lambu and Batu Punggul are each home to four begonia 
species (Kiew. 1998b) as, besides those listed in Table 1, B. gueritziana 
also grows there. 

While 17 species are narrow endemics, the eighteenth species, B. 
gueritziana, is not only widespread but is also the only species that is not 
confined to limestone. It grows on a variety of rock types usually by streams 
ranging from 30—750 m altitude, the latter on Gunung Kinabalu. However, 
although widespread, it has not been recorded from limestone in Sarawak. 
The distribution of begonias on limestone in Sarawak shows the same 
pattern of high endemism as in Sabah with species being restricted to 



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particular areas that also reflects the biodiversity patterns of other limestone 
species (Kiew, 1991), namely species are confined to the Bau area, the 
Subis area (Niah Cave) or to limestone in the Gunung Mulu National 
Park. The fact that only eight begonia species have been described from 
limestone in Sarawak, as opposed to 18 for Sabah, may be an indication of 
the lack of systematic botanical study than actual lower biodiversity. 

Of the 18 species now known from limestone in Sabah, 12 are new 
species described below. Apart from B. diwolii in sect. Diploclinium and 
the widespread B. gueritziana in sect. Platy centrum, the other 16 species all 
fall within sect. Petermannia. Section Petermannia is said by Doorenbos et 
al. (1998) to have male flowers typically with two tepals. However, as more 
species from Borneo are described, it is likely that this character will prove 
not to be diagnostic for the section as the majority of the cane-like begonias 
on limestone in fact have male flowers with four tepals. 

Section Petermannia is well represented in Borneo and includes 
several groups of similar species. One is the group of relatively short 
begonias recognised by Sands (1990) that have bristly stems, leaves that 
are narrow, obovate and with the midrib in line with the short petiole, the 
stipules persistent and becoming papery, and the flowers crowded into 
short inflorescences. From Sabah limestone, two species, B. anthonyi and 
B. berhamanii, belong to this group. These species do not grow directly on 
limestone but are common on the soil around the base of cliffs or on the 
steep slopes leading up to the hill, where they frequently form drifts. 

Another group includes the cane-like begonias that grow directly on 
the limestone. All these species are extremely decorative, particularly those 
with an unusual leaf shape (narrow and acutely pointed at both ends), with 
daintily scalloped leaf margins often coloured bright red, or with attractive 
variegation. In addition, B. keithii has striking red, glossy stems and the 
many small carmine heart-shaped male flower buds (Kiew, 1998c). Some 
can withstand full sun and live on the exposed summits, such as B. 
baturongensis, B. keithii, and B. madaiensis. These species do not flower in 
deep shade. Of the other species that grow in light shade beneath the tree 
canopy, B. amphioxus and B. layang-layang have narrow leaves with a 
pointed base, B. keeana, B. heliostrophe and B. malachosticta, have, in 
contrast, broader leaves with a large rounded basal lobe. These cane-like 
species have great potential as ornamental plants (Kee, 2000). Indeed, 
Sands (1990) reported that B. amphioxus grows well in hanging baskets. 

However, the affinity of some species in sect. Petermannia, such as 
B. lambii, B. gomantongensis Kiew, B. melikopia, B. postarii Kiew, B. 
punbatuensis and B. urunensis, is obscure. There is still a great deal to be 
learnt about the begonia flora of Borneo where the undescribed species 
outnumber those already named. 



Limestone Begonias of Sabah, Borneo 



245 



Key to Limestone Begonias in Sabah 

la. Small rhizomatous begonias with prostrate stems 2 

lb. Begonias with erect stems 3 

2a. Internodes up to 7.5 cm long, leaves well spaced: lamina margin 
hairy, fruit with 3 equal wings 5. B. diwolii 

2b. Internodes very short, leaves tufted; lamina margin glabrous, fruit 
with one wing longer than the other two 7. B. gueritizana 

3a. Stem short up to 13 cm tall with few (up to 4) leaves, lamina deeply 

corrugated 11. B. lambii 

3b. Stem more than 25 cm tall with many leaves, lamina plane 4 

4a. Mature stem conspicuously bristly, leaf with pinnate venation, basal 
lobe scarcely developed; male inflorescence compact, up to 2 cm 
long 5 

4b. Mature stem glabrous; leaf with palmate-pinnate venation, basal 
lobe well developed: male inflorescence a lax cymose panicle 7 

5a. Lamina lanceolate, up to 3 cm wide, basal lobe cuneate or slightly 
rounded, fruit up to 12 mm wide 4. B. berhamanii 

5b. Leaf obovate. 3 cm or more wide, basal lobe cordate or auriculate. 
fruit more than 14 mm wide 6 

6a. Leaf more than 2.5 times longer than wide, basal lobe cordate up to 
4 mm long, fruit wing tip pointed 2. B. anthonyi 

6b. Leaf up to 2.5 times longer than wide, basal lobe auriculate up to 10 
mm long, fruit wing tip rounded 18. B. urunensis 

7a. Lamina very hairy above 16. B. postarii 

7b. Lamina glabrous above 8 

8a. Lamina at least 4 times longer than wide, base narrowed to a point. 

sometimes peltate 9 

8b. Lamina less than 4 times longer than wide, base broad and rounded. 

never peltate 11 

9a. Lamina red-spotted: fruit stalk less than 2.5 mm long, fruit oblong 
up to 13 x 7 mm 1. B. amphioxus 

9b. Lamina plain green or grey mottled: fruit stalk more than 12 mm 
Ions, fruit ovoid or deltoid. 18 x 15 mm or larger 10 



246 Card Bull. Singapore 53 (2001 ) 



10a. Stem and petiole crimson; petiole up to 1.5 cm long; lamina not 
peltate; fruit ovoid and narrower, up to 25 mm wide .. 10. B. keithii 

10b. Stem and petiole pale green; petiole more than 2.5 cm long; lamina 

peltate; fruit deltoid and wider, more than 29 mm wide 

12. B. layang-layang 

11a. Lamina less than 6 cm wide 12 

lib. Lamina 6 cm or wider 14 

12a. Lamina more than 3 times longer than wide, pink spotted; tips of 
fruit wings pointed 14. B. malachosticta 

12b. Lamina less than 3 times longer than wide, not spotted; tips of fruit 
wings rounded 13 

13a. Petiole 3 — 4 cm long; fruit deltoid, shorter than wide, 22 — 28 mm 
wide 3. B. baturongensis 

13b. Petiole 1 — 3 cm long; fruit rhomboid (narrowed distally), longer 
than wide, 13 — 20 mm wide 13. B. madaiensis 

14a. Lamina more than twice as long as wide, apex pointing upwards ... 

8. B. heliostrophe 

14b. Lamina less than twice as long as wide, apex pointing downward... 

15 

15a. Lamina obliquely rotund; fruit oblong with pimply surface 

6. B. gomantongensis 

15b. Lamina obliquely ovate; fruit deltoid with smooth surface 16 

16a. Lamina less than 10 cm long; young stems minutely hispid; male 
flowers with outer 2 tepals 14 — 15 mm long 17. B. punbatuensis 

16b. Lamina more than 10 cm long; young stems glabrous; male flowers 
with outer tepals 7 — 8 mm long 17 

17a. Lamina up to 16 x 8.5 cm; petioles up to 7 cm long; female flowers 

solitary, male flowers with 4 tepals; fruit to 2.5 cm long 

9. B. keeana 

17b. Lamina 15 x 12 cm or larger; petiole more than 8 cm long; female 

flowers in pairs; male flowers with 2 tepals; fruit 4 — 5.5 cm long .... 

5. B. melikopia 



Limestone Begonias ofSabah. Borneo 



247 



1. Begonia amphioxus Sands 

Section Petermannia 

Sands. Kew Magazine. 7 (1990) 81 & Plate 149; Kiew, Gardenwise. 15 
(2000) 13. 

TYPE: Batu Punggul Sands 4045 (not seen). 

Distribution: Borneo - SABAH: Pensiangan District. Batu Punggul and 
Batu Tinahas. 

Habitat: Not common, base of limestone cliffs and in light shade on flat 
tops of subpeaks below the canopy. 

Notes: Begonia amphioxus is a striking begonia with reddish stems, peltate 
leaves narrowing sharply to a point at both ends, beautifully marked with 
deep crimson spots. It shows some variation in leaf shape (peltate or not), 
number of tepals in the female flower (3 to 5) and in the number of wings 
in the fruit (2 or 3). It is also unusual in having joined tepals in the female 
flower. 

Sands (1990) referred this species to sect. Platycentrum on account 
of the specimens he collected having female flowers with 2 locules and 
styles and 3 — 5 tepals. the male having 4 tepals. and the fruit being 2- 
winged although he noted that 'a third wing may occasionally develop in 
some fruits ... it tends to be smaller/ 

Collecting on Batu Punggul allowed the population at the type site 
to be reexamined as well as that on nearby Batu Tinahas. The majority of 
fruits on plants at the type site did indeed have fruits with two equal wings 
and two locules. but on the same plant, a few fruits with three wings (the 
third wing being smaller) could be found. On Batu Tinahas. plants mostly 
had fruits with three locules and three equal wings and only a few (on the 
same plant) had two locules and two wings. In all other respects the two 
populations were the same. The weak development of the third wing 
indicates a reduction from the typical state with three equal wings rather 
than the character state of sect. Platycentrum where the third wing is well 
developed and much larger. 

It is certainly an aberrant species for sect. Platycentrum. which, as 
Doorenbos et al. (1998) point out. is a section that includes plants that 
always have fruits with three wings, one of which is markedly larger than 
the other two. They drew attention to this anomalous situation by listing 
B. amphioxus under the heading "Species not attributable to any existing 
section'. As they noted, apart from the single character of the fruit being 
two locular, B. amphioxus is typical of sect. Petermannia in its erect habit, 
dichasial male inflorescences, solitary female flowers (which may have 4, 3 



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Card. Bull. Singapore 53 (2001) 



or 2 tepals but usually has 5) and the style, which is caducous in the fruit. 
On morphological grounds, there is therefore no doubt that B. amphioxus 
is a member of sect. Petennannia. In addition, recently Tebbitt (1999) 
using molecular data showed that it is affiliated with other species in sect. 
Petermannia and Tebbitt and Maciver (1999) demonstrated the presence 
of perforate base plates in its endothecial cells, again confirmation that it 
belongs to sect. Petermannia. 

Begonia amphioxus is easy to grow. In the Agricultural Park at 
Tenom, Sabah, it seeds freely and spreads on a lightly shaded limestone 
rockery. At Kew, it has been propagated from nodal cuttings, grows on all 
types of soil mix but best on a fibrous mix and it flourishes in hanging 
baskets (Sands, 1990). It certainly deserves to be more widely known in 
cultivation. 

Specimens examined: SABAH: Pensiangan District - Batu Punggul Ruth Kiew & S. 
Anthonysamy RK4379 (KEP, L, SAN, SING), Batu Tinahas Ruth Kiew & S. Anthony 
RK4337 (SAN, SAR, SING). 

2. Begonia anthonyi Kiew, sp. nov. 

Section Petermannia 

A Begonia cauliflora Sands foliis basi cordatis et tepalis integris glabris 
differt - - TYPUS: Batu Punggul R. Kiew & S. Anthonysamy RK 4352 
(holo SAN; iso K, SAR, SING). 

Figure 1 

Low, erect, unbranched herb, decumbent and rooting at nodes. Stem 
ferrugineous towards apex, 18 — 24 cm tall x 3 — 4( — 6) mm diam., internodes 
2 — 5.5 cm long, becoming woody, nodes not swollen. Stipules glabrous, 
narrowly lanceolate, 13 — 16 x c. 4 mm, margin entire, apex attenuate, 
persistent. Leaves alternate, distant, in life upstanding; petiole 2 — 4( — 15) 
mm long, slightly grooved above; lamina very dark plain green, papery 
when dry, glossy above, obovate, sometimes slightly falcate, (12 — )14( — 
17) x (3 — )4.5( — 6.2) cm, slightly asymmetric, narrow side curving inwards 
then rounded at base, broad side rounded (2 — )3( — 4) cm wide and cordate 
at base, basal lobes 2 — 4 mm long, margin dentate and sparsely fringed 
with bristly hairs, apex attenuate; midrib not at an angle to petiole, venation 
pinnate, lateral veins 6 — 7 pairs, branching towards margin, sometimes 
with 1 vein in basal lobe, impressed above and slightly prominent beneath. 
Plant protogynous. Male inflorescences from upper axils, erect, short 
flattened spike, 5 — 20 mm long of which peduncle is 5 — 7 mm, longer than 
petioles, continuously growing with one or sometimes two flowers open at 



Limestone Begonias of Sabah, Borneo 



249 




Figure 1. Begonia anthonyi Kiew 

A Habit x V,. B & C Male inflorescences x l'Aand x V 4 , D Male bud x V 4 . E Open male flower 
x 1 . F Androecium x 4, G Stamen x 8. H Female flower x 1,1 Open female flower x 1, J Style 
and stigma x 4. K T.S. ovary x 1V 2 , L Fruit x 7 4 . (from RK 4438) 



250 



Card. Bull. Singapore 53 (2001 ) 



a time; bracts distichous, pale green, persistent. Male flower with pedicel 
4 — 13 mm; tepals 2, totally white and scintillating, glabrous, slightly ovate, 
9 — 12 x 6 — 7 mm, margin entire, apex rounded; stamens c. 30, in a sessile, 
globose cluster c. 4 x 3 mm, filaments c. 1.25 mm long, anthers pale lemon 
yellow, ellipsoid, c. 1 mm long, apex emarginate. Female inflorescences 
with 1 or 2 flowers produced from lower leaf axils; bracts 2, reddish brown. 
Female flower with pedicel 4 — 6 mm, sparsely hispid; ovary pinkish with 
minute red hairs, narrowly deltoid, 9 — 16 x 6 — 16 mm, wings 3, equal, 
locules 3, placentas axile, bilamellate with many ovules on both surfaces; 
tepals 5, white or rosy red, elliptic, 6 — 10 mm long, outermost c. 6 mm 
wide, innermost c. 4 mm wide, margin entire, apex rounded; styles 3, c. 5 
mm long, joined c. half way then bifurcating; stigma papillose forming a 
continuous twisted band. Fruit pendent, pedicel stiff, 3 — 10 mm long, capsule 
broadly deltoid, 12 — 15 x 14 — 20 mm, locules 3, dehiscing between locules 
and wings, wings 3, 4 — 5 mm wide, thinly fibrous, tips pointed; seeds brown, 
minute, broadly ellipsoid, c. 0.25 mm long, base truncate, distally rounded. 

Distribution: Borneo: SAB AH - Pensiangan District, Batu Punggul, Batu 
Tinahas. 

Habitat: On rocks or on steep earth slopes up to the base of the limestone 
hill, often gregarious. 

Notes: Begonia anthonyi belongs to sect. Petermannia in being an erect, 
protogynous begonia with the ovary and fruit with three equal wings. Within 
sect. Petermannia, it conforms to the group of species recognised by Sands 
(1990) when he described B. cauliflora from Sabah that has hispid stems, 
short petioles, laminas that are widest at or above the middle and narrow 
to the base (rather than having a well developed basal lobe), has a midrib 
that is more or less in line with the petiole, and male flowers that are 
crowded in short axillary inflorescences. B. berhamanii described below 
also belongs to this group. In addition to the characters listed above, species 
in this group frequently have an unbranched stem, two tepals in the male 
flower, rather few stamens (to about 30), and the fruit stalk is thick and 
rigid. This last character contrasts with the other limestone species in this 
section where the pedicel is long and at fruit maturity is dry and thread- 
like so that the fruit is dangling. 

Begonia anthonyi and B. berhamanii grow on soil close to the cliff 
base or on the steep slope up to the hill. They tend to grow gregariously 
forming carpets but are local and not found away from the limestone hill. 

Begonia anthonyi is distinct from B. cauliflora in having a more 
asymmetric leaf with the broader side more rounded and it is cordate at 



Limestone Begonias ofSabah, Borneo 



251 



the base (not decurrent) and the lamina is green beneath (not crimson to 
brownish as in B. cauliflora), in the male tepals being white and glabrous 
(as opposed to pink and hirsute outside), and the female tepals glabrous 
and entire (hairy outside and serrate in B. cauliflora). In addition, in B. 
anthonyi flowers open one at a time, whereas B. cauliflora is shown as 
having several flowers open simultaneously on a single inflorescence. 

The species is named for S. Anthonysamy, for many years herbarium 
assistant at Universiti Pertanian Malaysia, who accompanied me and helped 
with the preparation of plant specimens on many field trips, including the 
one to Batu Punggul and Batu Tinahas. 

Specimens examined: SABAH: Pensiangan District - Batu Punggul the type and L. Kuntil 
SAN 135786 (SAN), Sumbing Jimpim SAN 136091 (SAN); Batu Tinahas R. Kiew & S. 
Anthonysamy RK 4438 (SAN, SING). 

3. Begonia baturongensis Kiew, sp. nov. 

Section Petermannia 

A Begonia keithii Kiew petiolis longioribus, foliis latioribus basi rotundatibus 
et fructibus latioribus differt — TYPUS: Batu Baturong R. Kiew et al. RK 
5026 (holo SAN; iso K, SAR, SING). 

Cane-like, glabrous begonia. Stem reddish-brown, glossy, erect, 30 — 50 cm 
tall and 5 — 10 mm thick, internodes (5 — )9.5 cm long, stems becoming 
horizontal with many leafy side shoots held horizontally, eventually falling 
and becoming pendulous, woody, slightly thicker and distinctly annular at 
nodes. Stipules pale green, slightly obovate to lanceolate, 25 — 28 x 10 — 11 
mm, entire, midrib keeled, apex acute, caducous. Leaves alternate, distant, 
pendant; petiole 3 — 4 cm, slender, slightly grooved above; lamina of young 
plants with light green or grey spots, adult unfolding leaves yellow becoming 
plain mid-green above and slightly flushed reddish-crimson, sometimes 
light green beneath, margin red, slightly succulent in live state, thinly 
leathery when dry, slightly glossy, obliquely ovate, asymmetric, (5.5 — )7 x 
4.25 — 5.5 cm, narrow side lanceolate, broad side broadly lanceolate 3.25 — 
4 cm long, base slightly cordate with basal lobe markedly rounded 2.5 — 4 
cm long, margin red, shallowly dentate, apex shortly acuminate; venation 
palmate-pinnate, midrib and lateral veins 4, ± equal-sized, branched c. 
midway to margin, c. 3 in basal lobe, prominent beneath. 
Plant protogynous. Male inflorescence axillary in distal leaf axils, glossy 
red, erect, cymose panicle to simple cyme in uppermost axil, longer than 
petioles, 3 — 4 cm long of which peduncle is 1.25 — 2 cm, much branched. 
Male flower with pedicel 2.5 — 5 mm long; tepals 4, outer surface of outer 2 



252 



Card. Bull. Singapore 53 (2001 ) 



cerise or deep rosy red, pinkish red inside, glabrous, outer two slightly 
cordate and concave, 6—7 x 6 — 8 mm, margin entire, apex acute, inner 
two pale pink, elliptic, 5 — 6 x 2.5 — 3.5 mm, apex acute; stamens more than 
30, cluster globose, sessile, 2.5 — 4 x 3 — 3.5 mm, filaments c. 0.5 — 0.75 mm, 
anthers obovate, c. 0.75 mm long, apex emarginate. Female flowers solitary, 
up to 5 each produced at successive axils, pedicel erect, 1 1 — 14 mm long, 
enclosed by 2 large bracts green tinged red, caducous; ovary green, obovate, 
16 — 20 x 12 — 17 mm, wings 3, equal, edges tinged red, locules 3, placentas 
axile, bilamellate with many ovules on both surfaces; tepals (4 — )5, rosy 
red, subrotund. slightly convex, isomorphic, 10 mm long, entire, apex 
rounded, outer 8 — 10 mm wide, innermost one c. 6 mm wide; styles 3, 
golden yellow, 4 — 5 mm long, free almost to base, bifurcating; stigma 
papillose forming a continuous twisted band. Fruit dangling on fine thread- 
like pedicel, 10 — 30 mm. capsule broadly ovate, (16 — )20( — 24) x (22 — ) 
24( — 28) mm, glabrous, locules 3, dehiscing between upper half of the 
locule and wing, wings 3 isomorphic, thinly leathery, truncate, c. 5 mm 
wide, thin and fibrous; seeds brown, minute, broadly ellipsoid, c. 0.3 mm 
long, base truncate, rounded distally. 

Distribution: Borneo: SABAH - Lahad Datu District. Bukit Baturong. 

Habitat: Begonia baturongensis grows on limestone rock below the canopy 
in light shade or on exposed rock ledges and crags. 

Notes: This species is typical of sect. Petermannia in its cane-like habit and 
the ovary and fruits having three equal wings. In addition, the female 
flowers produced in the lower leaf axils with the male inflorescences in the 
upper ones is a condition found in species of this section. 

Like Begonia madaiensis (described below), B. baturongensis has 
relatively small leaves, which decrease markedly in size towards the apex, 
which are well-spaced on long petioles with their apex pointing downwards 
resulting in the twigs having a dainty zigzag appearance, especially on the 
lateral horizontal branches. The juvenile leaves of both are variegated, but 
become plain green with age. The strikingly variegated juvenile leaves are 
illustrated by Kiew (2000). Male inflorescences are only produced after the 
female flowers have already become fruits. 

In addition, these two species share the same niche growing fully 
exposed or in light shade on the summit and shoulders of cliff faces and, 
indeed, they do not flower in shade. 

The two species have very different fruits: those of B. baturongensis 
are broader than long and have truncate wing tips, while those of B. 
madaiensis are longer than wide and are narrowed distally. The leaves are 



Limestone Begonias of Sabah. Borneo 



253 



also different, those of B. baturongensis have longer petioles (3 — 4 cm 
long) and the leaf margin is shallowly dentate, while the leaves of B. 
madaiensis have short petioles (up to 3 cm long) and the margin is scalloped. 
The flowers of B. baturongensis are more decorative being deep rosy red 
to cerise compared with the white flowers of B. madaiensis. 

Specimens examined: SABAH: Lahad Datu District - Bukit Baturong: the tvpe and S.P. 
Lim et al. LSP 737 (SAN, SING). 

4. Begonia berhamanii Kiew, sp. nov. 

Section Petermannia 

A Begonia cauliflora Sands foliis angustioribus et tepalis integris glabris 
differt — TYPUS: Batu Punggul R. Kiew & S. Anthonysamy RK 5046 
(holo SAN; iso SING). 

Figure 2 

Low, erect herb, becoming procumbent and rooting at nodes, indumentum 
of short hooked hairs, appressed and dense on stem, petiole and lower 
surface of veins. Stem reddish-brown, up to 25 cm tall and 2 — 3 mm diam., 
unbranched, woody, internodes 1.5 — 2 cm long, nodes swollen. Stipules 
lanceolate. 13 — 14 x 6 — 7 mm, midrib dorsally strongly keeled with minute 
scattered hairs, margin entire with a row of fine hairs, apex narrowly 
attenuate, persistent. Leaves alternate, distant, held horizontally: petiole 
3 — 5 mm long, shallowly grooved above: lamina plain dark green and 
glabrous above, whitish green (sometimes faintly reddish) beneath, papery 
when dry. glossy above, slightly asymmetric, narrowly lanceolate and slightly 
falcate, 12.5 — 16 x 2.5 — 3.2 cm of which broad side 1.5—1.8 cm wide, basal 
lobe cuneate sometimes rounded, 2 — 3 mm long, margin shallowly dentate, 
apex attenuate; venation pinnate, midrib and lateral veins concolorous with 
lamina. 7 — 8 pairs, branching c. midway to margin, 1 vein in basal lobe, 
slightly impressed above, slightly prominent beneath. 
Plant protogynous. Male inflorescences from upper leaf axils, sessile, to 2 
cm long, shorter than petioles, sometimes bifurcating, flowers congested 
and covered by distichous, overlapping bracts, only one or two flowers 
open at a time: bracts deep rosy or brown red, lanceolate, c. 12 x 3.5 — 4 
mm. margin dentate with fringe of hairs, persistent. Male flower with 
minutely hairy, pale pink pedicel 4 — 11 mm long, tepals 2, opening rosy 
red, fading to white faintly tinged pink outside, elliptic, 6 — 8 x 6 — 7 mm, 
margin entire, apex rounded; stamens 20 — 30, in hemispherical sessile cluster 
c. 3 mm across, filaments 0.5 — 1.5 mm long, anthers dull white, rotund, c. 
0.75 mm long, apex emarginate. Female flowers solitary from lower leaf 



254 



Card. Bull. Singapore 53 (2001 ) 



axils; pedicel 2 — 3 mm long, with densely appressed hairs; ovary white, 
deltoid, 8 — 12 x 11 — 16 mm, locules 3, outer surface minutely hairy, wings 
3, equal, 4 — 5 mm wide, glabrous, placentas axile, bilamellate with many 
ovules on both surfaces; tepals 5, rosy red, broadly elliptic, isomorphic, 5 — 
10 mm long, apex rounded, outer 4.5 — 6 mm wide, inner 3 — 4.5 mm wide, 
margin entire; styles 3, 4 — 6 mm long, divided to base, bifurcating; stigma 
papillose forming a continuous twisted band. Fruit with stiff, decurved 
pedicel c. 3 — 4 mm long, the fruit becoming bent backwards and parallel 
to stem, capsule broadly deltoid, 11 — 13 x 10 — 16 mm, sparsely hispid, 3- 
locular, dehiscing between locule and wing, wings 3, isomorphic, 2.5 — 8 
mm wide, tips slightly acute, thin almost papery; seeds brown, minute, 
almost globose, c. 0.2 mm long, narrowed to base, rounded distally. 

Distribution: Borneo: SAB AH - Lahad Datu, Bukit Baturong. 

Habitat: Shaded base of limestone cliff on damp soil. 

Notes: Like Begonia anthonyi, this species belongs to the group of species 
within sect. Petermannia with hispid stems, short petioles, slightly 
asymmetric laminas with a poorly developed basal lobe and with the midrib 
almost in line with the petiole. B. berhamanii more resembles B. cauliflora 
Sands in its narrowly lanceolate leaves with a slightly rounded basal lobe 
and in its dense clusters of male flowers. It is readily distinguished from B. 
cauliflora by its narrower leaves with an attenuate apex that are whitish 
green beneath (in B. cauliflora the leaves are 4.2 — 6.4 cm wide, the apex is 
acute-acuminate and they are crimson to brownish beneath) and the 
glabrous, entire tepals of the male and female flowers (those of B. cauliflora 
are hairy outside and those of the female flower are serrate). 

The species is named for Berhaman Ahmad who, while he was a 
research officer at the Forest Research Centre, Forest Department Sabah, 
ably organised several expeditions to explore and collect the limestone 
flora. 

Specimens examined: SABAH: Lahad Datu District - Bukit Baturong the type and S.P. 
Lim et al. LSP 715 (SAN, SING). 

5. Begonia diwolii Kiew, sp. nov. 

Section Diploclinium 

A Begonia speluncae Ridl. caulibus pilosis, internodiis longioribus et foliis 
non-peltatis differt — TYPUS: Tempadong, Segama River Ruth Kiew et 
al. RK 4767 (holo SAN, iso BRUN, K, L, SAR, SING). 




Figure 2. Begonia berhamanii Kiew 

A Habit x V : ,B Stipules x 2. C Male flowers x 2. D Fruit x 1 7 : . (from LSP 715) 



256 



Curd. Bull. Singapore 53 (2001) 



Figure 3 

Creeping, prostrate herb attached to substrate at each node by fibrous 
roots, old stems setose, indumentum of appressed, concolorous, uniseriate, 
eglandular hairs c. 1.5 mm long on the young stems, stipules, petiole, leaf 
margin, lower surface of lamina and particularly dense on veins. Stem pale 
purplish becoming deep reddish brown with age, up to c. 1 m long and 
1.5 — 2 mm diam. near apex and to 4 mm in old stems, succulent, not 
swollen at nodes, internodes 4 — 12 mm apart at apex lengthening to 5.5 — 
7.5 cm, stem slightly zigzag with short lateral prostrate branches, c. 10 cm 
long produced at wide intervals. Stipules pale reddish purple, broadly 
lanceolate, 9 — 19 x 3.5 — 8 mm, margin entire, apex distinctly setose, setae 
4 — 5 mm long, persistent. Leaves alternate, distant, lamina positioned flat 
against rock surface; petiole brownish or reddish purple, more hispid than 
stem, 3 — 7 cm long up to 14.5 cm on pendent branches, 1 — 3 mm diam.. 
terete; lamina rich deep plain green above, deep magenta beneath, thinly 
fleshy, matt, upper surface glabrous except for a few setose hairs on the 
veins, orbicular to reniform, smaller leaves slightly asymmetric, large leaves 
very asymmetric, (2 — )4( — 6.5) cm long and (2.5 — )5( — 6.5) cm wide of 
which the broader side is (1.5 — )3( — 4.5) cm wide, base slightly cordate 
but not overlapping, basal lobes shallowly rounded (0.5 — )1.5( — 3.5) mm 
long, margin slightly wavy and fringed by long hairs, apex rounded in 
orbicular leaves, acute in reniform leaves; venation palmate, midrib and 
lateral veins concolorous with lamina (occasionally paler), (2 — )3, equal 
sized, bifurcating twice, first near petiole or more than half way towards 
margin, second close to margin, 1 — 2 veins in basal lobe, slightly impressed 
above, slightly prominent beneath. 

Inflorescence protandrous, axillary from second node, cymose panicle, pale 
magenta, sparsely setose, erect, longer than petioles, 7 — 15 cm long, 
peduncle 6.5 — 10.5 cm long, each branch with 1 female flower and several 
male flowers eventually with up to 3 branches; bracts paired at base, brown 
and papery, narrowly acute with conspicuous midrib, 6 — 9 x 2.5 — 3 mm, 
margin entire, apex setose, caducous. Male flowers with pale pink pedicel 
7 — 10 mm long; tepals 4, outer 2 with the upper tepal deep rosy pink 
outside, both suffused pink inside, glabrous, broadly oval, 6 — 9 x 5 — 7 mm, 
margin entire, apex rounded to acute, inner 2 almost completely white, 
oval, narrower, 7 — 8 x (2.5 — )4 mm, apex bluntly rounded; stamens c. 30, 
cluster subglobose with torus c. 1 mm long, filaments c. 0.4 mm long, 
anthers lemon yellow, obovate, c. 0.6 x 0.5 mm, apex emarginate. Female 
flower with pale pink to magenta pedicel c. 6 mm long; ovary pale pink 
with margin of wings tinged green, sparsely pustulate with occasional short 
hairs, cordate, c. 3.5 x 3.5 mm, wings 3, equal-sized, c. 1.25 mm wide, base 



Limestone Begonias ofSabah. Borneo 



257 




Figure 3. Begonia diwolii Kiew 

A Habit x'A.B Male flower. C Open male flower. D Androecium. E Stamen. F Female flow. 
G Style and stigma. H T.S. ovary. I Fruit, (from RK 5073) 



258 



Card. Bull. Singapore 53 (2001) 



rounded, narrowed to apex, locules 3, placentas axile, bilamellate with 
many ovules on both surfaces; tepals 4, c. 4.5 — 8 x 4 — 5 mm, margin entire, 
outer 2 slightly smaller, tinged magenta with outer surface paler inside, 
scintillating, microscopically pustulate, apex rounded; inner 2 slightly longer, 
pure white, apex truncate; styles 3, c. 3.5 mm long, joined at the base for c. 
1.5 mm, then bifurcating; stigmas green-yellow, with a lunate, papillose 
band on each branch. Fruits up to 3 per infructescence, pendent, peduncle, 
branches and pedicels becoming thin, dry and hair-like, 12.5 — 16.5 cm 
long, capsule 10 — 13 x 3.5 — 5 mm, whole fruit becoming dry and papery, 
glabrous, locules 3, dehiscing between locule and wing, wings 3, equal- 
sized, broadly rounded, c. 4 — 5 mm wide; seeds brown, minute, broadly 
ellipsoid, c 0.25 — 0.3 mm long, base narrowed, apex rounded. 

Distribution: Borneo - SABAH: Lahad Datu District, Segama River 
(Tempadong, Batu Belas, Borneo Marble Quarry and an unnamed hill by 
the Lahad Datu road). 

Habitat: On vertical and horizontal rock faces from the cliff base in deep 
shade to light shade near the summit of limestone hills. 

Notes: This new species is most distinctive in its habit of producing long 
prostrate, frequently zigzag stems that creep and root over the rock surfaces 
(illustrated in Kiew, 2000). The long internodes mean that the leaves are 
distant and so contrast with other rhizomatous limestone species with leaves 
with palmate venation, such as B. gueritziana in Sabah and B. speluncae 
Ridl. in Sarawak, which both have very short internodes so that their 
leaves are in a tuft. Furthermore, it cannot be mistaken for B. gueritziana, 
which has much larger and thicker leaves, and fruits with one wing much 
larger than the other two. In fruit shape (three equal, narrow, rounded 
wings), leaf shape (small and frequently orbicular) and the inflorescence 
being longer than the petiole, B. diwolii most closely resembles B. speluncae 
but it is readily distinguished from the latter, which not only has short 
internodes and tufted leaves, but is also glabrous, and the leaves are 
frequently peltate. In addition, B. speluncae is placed in sect. Reichenheimia 
by Doorenbos et al. (1998), a section that has placentas with one (not two) 
lamellae. 

This new species belongs to sect. Diploclinium Group 1 of Doorenbos 
et al. (1998), which includes the rhizomatous species with palmately veined, 
often symmetric leaves, bisexual inflorescences that are usually protandrous, 
male flowers with four tepals and obovate anthers, female flowers with 
four tepals, and three-loculate fruits with three equal-sized wings and 
bilamellate placentas. 



Limestone Begonias ofSabah, Borneo 



259 



Many of these characters are shared by sect. Petermannia. although 
species in that section are less usually rhizomatous or palmately veined. 
However, the one character that does exclude B. diwolii from sect. 
Petermannia and places it in sect. Diploclinium is its protandry. 

The fact that this species is protandrous is not always apparent from 
herbarium specimens. Observations in the field show that the inflorescences 
produce three branches in sequence, each of which produces one or two 
male flowers that open first, followed by a single female flower, then further 
male flowers, the entire inflorescence therefore eventually producing a 
maximum of three fruits. Unless a plant is observed in the initial male 
phase, it appears protogynous with a developing fruit below and young 
male flowers above. 

This charming species with its dainty round leaves magenta beneath 
and bright pale flowers deserves to be introduced into cultivation. 
Experimentation in the Singapore Botanic Gardens shows that it grows 
well on old coral (Kee, 2000). 

It gives me great pleasure to name this decorative species for Diwol 
Sundaling. Senior Herbarium Officer at SAN, who organised the expedition 
to the Segama River (and many others) and whose enthusiasm and support 
in the field is much appreciated. 

Specimens examined: SABAH: Lahad Datu District - Tempadong the type and Ruth Kiew 
et al. RK5073 Unnamed hill on Lahad Datu road (E, SAN, SING). 

6. Begonia gomantongensis Kiew 

Section Petermannia 

Kiew. Gardens" Bulletin Singapore 50 (1998) 164. 

TYPE: Bukit Dulang Lambu James Awing SAN 47257 (holo SAN). 

Distribution: Borneo: SABAH - Kinabatangan District. Bukit Dulong 
Lambu (Gomantong Cave). 

Habitat: Base of limestone hill in deep shade on boulders or the foot of 
cliff faces. 

Notes: A member of sect. Petermannia, it is distinct from the other limestone 
species in Sabah in its subrotund leaves, which are held horizontally on 
long petioles, and its oblong capsules with a pimply surface. 
It is known only from Bukit Dulong Lambu. 

Specimens examined: SABAH: Kinabatangan District - Bukit Dulong Lambu James Awing 



260 



Card. Bull. Singapore 53 (2001) 



SAN 47257 (SAN;, S.P. Lim & Ubaldus LSP 785 (SAN, SING), R. Kiew & S.P. Lim BDL 
3 (SAN, SING). 

7. Begonia gueritziana Gibbs 

Section Plaiy centrum 

Gibbs, Linn. Soc. Bot. 42 (1914) 82 & figure. 

TYPE: Kayoh Hills, Tenom L.S. Gibbs 2892 (holo BM). 

As more specimens are available, including ones with female flowers, 
additional information is provided here to supplement the original 
description: 

Rhizome with crowded internodes. Petiole 6 — 13 cm long; lamina dark 
green above, often reddish-purple beneath, 5 — 8 x 5.5—7 cm, basal lobes 
scarcely developed 3—18 mm long. Inflorescences protandrous, longer than 
petiole, 11 — 22 cm long of which peduncle is 10 — 20 cm. Male flower outer 
two tepals broadly elliptic, 11 — 12 x 7.5 — 8 mm, inner two narrowly elliptic, 
c. 11 x 3 mm; stamens joined at base, torus c. 1 mm long. Female flowers 
with ovary pale green, glabrous, c. 5 — 9 mm long, locules 2, placentas 
axile, bilamellate, wing 3 unequal, longer wing c. 4 — 7 mm wide, two shorter 
c. 2 mm wide; tepals 4, rosy pink, outer two almost rotund, 5 — 9 x 5 — 8 
mm, margin entire, apex rounded, inner two elliptic, c. 4 — 8 x 4 — 5 mm, 
apex cucullate; styles yellow, 2.5 — 3 mm long, bifurcating; stigma a twisted 
continuous papillose band. Capsule c. 10 mm long, longer wing 10 — 12 mm 
wide, broadly rounded, slightly concave, two shorter wings 4 — 5 mm wide, 
thin, apex slightly acute. 

Distribution: Borneo - SABAH: Apart from islands and the coastal areas 
of the west and north coasts, B. gueritziana is widespread in Sabah both on 
and off limestone having been collected from the Tenom. Keningau, Kota 
Merudu, Ranau, Labuk Sugut, Kinabatangan and Tawau districts. However, 
it has not been collected further west than the Kallang Waterfall in Tenom 
(Ruth Kiew & S.P. Lim RK 4280). It has not been reported from Brunei 
(Sands, 1996) nor has it been collected from Gunung Api, in the Mulu 
National Park. Sarawak, the closest limestone to Sabah. It therefore is a 
Sabah endemic. 

It is the only widespread begonia species on limestone in Sabah and 
is particularly common on all the Kinabatangan limestone hills (even on 
the smaller, disturbed ones like Supu and Batu Tulug) and has been 
collected from the Kelabangan Ridge, Batu Urun and Lian Cave. On the 
other hand, it is absent from limestone in the south (Segama River and 



Limestone Begonias of Sabah, Borneo 



261 



Madai-Baturong limestone) and in the west (Melikop, Pun Batu, Batu 
Punggul and Sungai Pangi). 

Habitat: On limestone, B. gueritziana grows in light shade and is most 
frequent in soil-filled crevices and is particularly common on humus covered 
ledges (Kiew, 1998b). 

Of the begonia species that grow on limestone in Sabah, it is the only 
species that has been collected from non-limestone substrates (basalt, 
gabbro and sandstone are cited on herbarium specimens). It has usually 
been collected from rocks by the edge of streams from near sea level (30 
m) up to 750 m altitude on Gunung Kinabalu. 

Notes: Gibbs (1914) originally placed B. gueritziana in sect. Reichenheimia 
as do Doorenbos et al. (1998). However, it is atypical of this section in the 
ovary having unequal wings, being 2-loculate and each locule having two 
placentas. These ovary characters clearly place it within sect. Platy centrum, 
with which it is in accordance for all other diagnostic characters, such as 
the rhizomatous habit, palmate leaf venation, the axillary, protandrous 
inflorescence (illustrated in Kiew, 2000), male flower with 4 tepals, and the 
nodding fruit, which dehisces on both sides of the narrower wings. 

As in B. diwolii, examination of herbarium specimens can be 
misleading, because if the initial stage with only male flowers is not 
represented, the inflorescence may appear to be protogynous. 

Compared with the begonia flora of Peninsular Malaysia, sect. 
Platycentrum is poorly represented in Sabah, the only other species being 
B. adenostegia Stapf from Gunung Kinabalu. 

Although widespread in terms of geography, substrate and altitude, 
it is rather uniform in habit and leaf shape. However, the population on 
limestone at Kelabangan is different in having pure white flowers compared 
to the usual rosy pink ones. 

It is apparently an easy species to grow, as at the Agricultural Park. 
Tenom, it seeds freely and has gone wild on shaded rocks in the Orchid 
Centre. 

Specimens examined: The type and from limestone: SABAH: Kinabatangan District - 
Kinabatangan Valley. Batu Batangan Ruth Kiew & S.P. Lim RK 4291 (SAN, SING). 
Meijer SAN 23103 (locality misnamed Batu Bilit) (SAN); Batu Temambong Besar Ruth 
Kiew & S.P. Lim RK 4192 (SAN, SING), Lim el al. LSP 1151 (SAN, SING); Keruak FR 
Ruth Kiew & S.P. Lim RK 4183 (SAN, SING); Bukit Dulong Lambu (Gomantong) Meijer 
SAN 20754 (K), S.P. Lim et al., SPL 616 (SAN, SING); Supu /. Singh & Eging SAN 51826 
(SAN). Batu Tulug (Batu Putih) Dewol & Harun SAN 89912 (SAN, SING), Puasa SAN 
10115 (K). S.P. Lim et al. LSP 625 (SAN. SING). LSP 771 (SAN. SING); Pensiangan 
District - Batu Urun Ruth Kiew & S. Anthony RK4463 (SAN, SING) - also observed on 



262 



Gard. Bull. Singapore 53 (2001) 



Baladut and Sarupi (Kinabatangan Valley); Kelabangan (Pensiangan District). 

8. Begonia heliostrophe Kiew, sp. nov. 

Section Petermannia 

A Begonia malachosticta Sands foliis majoribus sursum spectantibus differt 
— TYPUS: Batu Batangan R. Kiew & Lim S.P. RK 4293 (holo SAN; iso 
BRUN, K, KEP, L. SAR, SING). 

Figure 4 

Bushy, cane-like, glabrous begonia, usually branched at base from a 
prostrate rhizome. Stem reddish brown or purple, up to 1.25 m x c. 7.5 mm 
diam., becoming woody, flowering at 60 cm, internodes 5 — 7 cm long, 
erect, branching at c. 30 cm, nodes swollen and almost articulate. Stipules 
pale green, lanceolate, 10 — 15 x 4 — 6 mm, margin entire, apex setose, 
caducous. Leaves alternate, distant, positioned with the apex held upwards; 
petiole reddish brown or purple, 5 — 7 cm long, grooved above; lamina 
sometimes with a silvery hue when immature, becoming flecked silver and 
finally uniformly mid-green at maturity, except for deep crimson patch at 
junction with petiole, beneath slightly reddish-green between veins, thinly 
succulent in life, thinly chartaceous dried, glossy above, asymmetric, 
narrowly ovate (10.5 — )14 — 16( — 19) x (3.5 — )6 — 6.5 cm of which broad 
side is (2.5 — )4.5 — 5 cm wide, basal lobe rounded, (2 — )3.5 — 5 cm long, 
base cordate but not overlapping, margin reddish, serrulate, apex attenuate; 
venation palmate-pinnate, midrib and lateral veins 3 — 5 pairs along midrib 
and 1 pair at base, bifurcating once about halfway to margin and again 
close to the margin, 2 — 3( — 4) veins in basal lobe, slightly raised above, 
plane and pale green beneath. 

Inflorescences axillary, protogynous, much branched cymose panicle, bright 
deep magenta, erect, 7.5 — 10 cm long of which peduncle is 1.75 — 3.5 cm 
long, longer than petioles, with 1 or 2 female flowers proximally and many 
male flowers distally on thin, branches; bracts lanceolate, pale green or 
reddish green, c. 7 x 4 mm, margin entire, apex acute, caducous. Male 
flower with reddish pedicel c. 5 mm long; tepals 4, white or very pale green, 
suffused crimson towards base, inner surface glistening, margin entire, apex 
rounded, outer two concave, almost rotund 4 — 6 x 4.5 — 6 mm, inner two 
broadly elliptic 2 — 4 x 1.5 mm; stamens c. 60, cluster broadly conical, c. 3 
mm long including a short torus c. 0.25 mm long, filaments c. 0.75 mm long, 
anthers lemon yellow, broadly oblong, c. 0.5 mm long, apex emarginate. 
Female flower pendent in bud then straightening so the open flower is held 
horizontally; pedicel 1.75 — 2 cm long, crimson, ovary glabrous, oblong, c. 



Limestone Begonias o f Sabah. Borneo 



263 




Figure 4. Begonia hehostrophe Kiew 

A Habit x 1 A. B Branchlet of male inflorescence x 2. C Male bud x 2. D Open male flower x 2 V 2 , E 
Androecium x 5, F Stamen x 10. G 1&2 Female flower x 1. H Open female flower x 1V 2 , 1 
Style and stigma x 5. J T.S. ovary x 1. (from RK 4293) 



264 



Card. Bull. Singapore 53 (2001) 



15 x 9 mm wide proximally and 14 mm wide distally, wings 3, isomorphic, 
reddish when immature becoming green with slight reddish tinge on the 
margins at maturity, locules 3, placentas axile, bilamellate, many ovules on 
both surfaces; tepals 5( — 6), pale yellowish green with a peachy tinge, 
elliptic, isomorphic, c. 10 mm long, outermost 6 — 9 mm wide, innermost c. 
4 mm wide, margin entire, apex rounded; styles 3, c. 3 mm long, divided to 
base, bifurcating, stigma bright lemon yellow, papillose forming a continuous 
twisted band. Fruit dangling on a fine hair-like pediceh 2 — 3 cm long; 
capsule broadly deltoid, 17 — 23 x 17 — 22 mm, glabrous, locules 3, dehiscing 
between locule and wing, wings isomorphic, rounded at base, 7 — 10 mm 
wide, tips rounded, thinly fibrous; seeds brown, minute, broadly ellipsoid, 
c. 0.25 — 0.3 mm long, base truncate, rounded distally. 

Distribution: SABAH: Kinabatangan District, Kinabatangan Valley (Batu 
Batangan and Keruak). 

Habitat: On soil at the base of cliffs or in crevices in vertical rock faces 
from the base to the crest of cliffs in deep or light shade, particularly 
common on soil-covered ledges in light shade. 

Notes: In its cane-like habit, its female flowers with 5 tepals, 3 bifurcating 
styles that are caducous in the fruit, 3-loculate ovary and fruit with 3 equal 
wings and bilamellate placentas, it is typical of sect. Petermannia but like 
the majority of limestone begonias, it differs in having 4 tepals in the male 
flower instead of the typical 2. 

It is unique among the limestone begonias in Sabah in that the leaf 
grows with its apex pointing upwards compared with the usual position 
where the apex points downward. The species epithet highlights this unusual 
feature. 

It belongs to a group of cane-like begonias that grow in light shade 
on rock faces that have asymmetric, narrowly elliptic leaves with one basal 
lobe well-developed and rounded. On limestone, this group includes 
Begonia tnalachosticta, B. keeana and the above species. In addition to the 
orientation of the leaf, B. heliostrophe differs from B. malachosticta in its 
broader, plain (non-variegated leaves), many-flowered male inflorescences 
and rounded (non-arcuate) wings of the fruit, and from B. keeana, besides 
the orientation and non-variegation of the leaf, by its larger leaves (up to 
12.5 x 8.5 cm in B. keeana) and yellowish green flowers (compared with 
the rosy red ones of B. keeana). 

Specimens examined: SABAH: Kinabatangan District - Batu Batangan the type; Keruak 
Amin & Matin SAN 108091 (SAN), Ruth Kiew & Lim S.P. RK 4158 (E, K, L. SAN, SAR. 
SING), RK 4175 (SAN, SING). 



Limestone Begonias ofSabah, Borneo 



265 



9. Begonia keeana Kieu, sp. nov. 

Section Petermannia 

A Begonia malachosticta Sands foliis latioribus et fructibus latioribus 
pedicello multo longiore suffultis — TYPUS: Tempadong R. Kiew et al. 
RK 4766 (holo SAN: iso K. SING). 

Cane-like begonia, glabrous, root stock woody with several stems. Stem 
reddish brown flecked white, smooth and glossy, to 1.25 m tall and 5 — 7 
mm diam.. flowering at c. 30 cm tall, internodes 2 — 8.5 cm. erect or pendent 
if growing on edge of cliff face, older stems branching sparingly in the 
upper 50 cm. branches at 45' to main stem, woody, slightly swollen at 
nodes. Stipules pale green, broadly lanceolate, 14 — 23 x 7 — 10 mm, margin 
entire, apex setose, persistent. Leaves alternate, distant, lamina pendent 
held at 45 on erect petioles: petiole concolorous with stem, in lower leaves 
c. 3 — 7 cm x 3 — 4 mm long, in upper leaves 1 — 1.5 cm x c. 2 mm long, 
terete: lamina variegated, margin silvery grey, veins dark green, lamina 
between veins either completely silvery grey to the base of the veins or 
with a few. large grey-green blotches, beneath uniform deep purple magenta, 
fleshy (brittle and snapping in live state), thinly chartaceous in dried state, 
glossy above, obliquely narrowly ovate, asymmetric. 11.5 — 16 x 7—8.5 cm, 
the broader side 5 — 5.5 cm wide, base cordate but not overlapping, basal 
lobes rounded. 4 — 5 cm long, margin slightly scalloped between veins and 
minutely serrulate, apex attenuate: venation palmate-pinnate, midrib and 
lateral veins 3 — 4, equal-sized, branching midway to margin, 2 veins in the 
basal lobe, slightly indented above, slightly prominent and concolorous 
beneath. 

Plant protogynous. Male inflorescence produced after and in the same axil 
as a fruit or from the upper leaf axils, an erect, cymose panicle with third 
order branching, reddish brown or deep purple red, longer than subtending 
petiole. 4 — 15.5 cm long of which peduncle is 2.5—7.5 cm long: bracts pale 
green, ovoid. 6 — 9 x 5 — 6 mm. margin entire, apex acute, persistent. Male 
flowers with pedicel rosy red. 13 — 18 mm in open flower, very slender. 
tepals 4. pale rosy red deeper red towards centre, margin entire, outer two 
± rotund, cucullate. 7 — 8 x 7 — 9 mm. glabrous, inner two oblong, apex 
rounded, 6 — 8 x 2 — 3.5 mm: stamens more than 40, cluster hemispherical, 
3( — 4) x 3( — 4) mm including torus c. 0.5 mm long, filaments c. 0.75 — 1 
mm long, anthers pale yellow. 0.75 — 1 mm long, apex emarginate. Female 
flowers solitary, produced at 1 or 2 ( — 3 — 4) successive leaf axils: pedicel c. 
15 — 16 mm long; ovary deep rosy red. oblong, 16 — 24 x 15 — 22 mm, wings 
3. equal, locules 3. paler, placentas axile, bilamellate with many ovules on 
both surfaces: tepals 5, outermost glossy, rosy red (paler than wings), broadly 



266 



Card. Bull. Singapore 53 (2001) 



oval c. 9 — 10 x 8 — 9 mm, innermost isomorphic and slightly smaller c. 7 x 6 
mm. sometimes narrowly elliptic c. 8 x 4 mm. margin entire, apex rounded: 
styles 3, pale yellow, c. 4 mm long, joined at base for c. 1.5 mm, bifurcating: 
stigma papillose forming a continuous twisted band. Fruit dangling, pedicel 
drying to fine thread, 23 — 30 mm long, capsule broadly deltoid, 25 — 31 x 
25 — 29 mm, glabrous, locules 3, dehiscing on upper half between locules 
and wing, wings 3. equal, 13 — 15 mm wide, very thin and fibrous, tip 
rounded; seeds brown, minute, ellipsoid, c. 0.3 mm long, truncate at base, 
rounded distally. 

Distribution: Borneo - SABAH: Lahad Datu District. Tempadong and 
Batu Belas on Segama River and Tabin Wildlife Reserve. 

Habitat: At base of cliffs, in crevices in rock face or on exposed shoulders 
of cliffs below tree canopy in light shade. 

Notes: It is a typical member of sect. Petermannia in its erect canedike 
habit, asymmetric leaves, in being protogynous with solitary female flowers 
produced before the distal, many flowered male inflorescences, in having a 
3-loculate ovary with the placentation axile. bilamellate placentas, and the 
fruit having three equal wings and caducous style. 

It most resembles B. malachosticta Sands in its habit and leaf shape 
but differs in the pattern of variegation (B. malachosticta has pink spotted 
leaves), much wider leaves (they are less than 5 cm wide in B. malachosticta). 
much branched male inflorescence (that of B. malachosticta is a simple 
cyme with three flowers or is once branched and just 4 cm long with about 
six flowers), larger male flowers (3 — 7.5 mm long in B. malachosticta) and 
the broader fruit with rounded wings (fruit is narrower. 10 — 17 mm wide, 
and has arcuate wings in B. malachosticta). 

Local people eat the young, succulent shoots of this new species as a 
vegetable. Most begonia leaves have a sourish taste, but in Sabah relatively 
few are actively gathered for food. Another such esculent species. Begonia 
lazat Kiew & Reza Azmi, was discovered during the course of an 
ethnobotanical study (Reza Azmi and Kiew. 1998). The fact that species 
known to local people as food plants are still unknown to science indicates 
how little is still known about the begonia flora in Sabah. 

This is a handsome begonia illustrated in Kee (2000) and Kiew (2000). 
which has lush, glossy leaves with silvery variegations on the upper surface 
and a deep magenta underside. It grows well on old porous coral in the 
nursery at Singapore Botanic Garden (Kee, 2000). That it adopts a 
pendulous habit when growing on the edge of cliffs indicates its potential 
for being grown in hanging baskets. 



Limestone Begonias ofSabah. Borneo 



This new species is named for Andrea Kee. horticulturalist at 
Singapore Botanic Gardens, who was a member of the expedition that 
discovered this new species and who has carried out trials on its cultivation. 

Specimens examined: SABAH: Lahad Datu District - Segama River - Batu Belas Ruth 
Kiew et al. RK 4765 (SAN. SING): Tempadong the type: Tabin Wildlife Reserve Ruth 
Kiew RK 5118 (SAN. SING. University of Sabah). 

10. Begonia keithii Kiev* 

Section Petermannia 

Kiew. Gardens" Bulletin Singapore 50 (1998) 189 & Fig. 4. 

TYPE: Batu Tengar Cave Kiew, Anthony & Urn RK4327 (holo SING: iso 

K. KEP. L. SAN. SAR 

Figure 5 

Distribution: Borneo - SABAH: Semporna District. Batu Tengar Cave 
(Segarong Cave). 

Habitat: In light shade or full sun. growing in crevices on the limestone hill, 
dominating exposed shoulders and summit of the hill, where it forms thickets 
with its woody cane-like stems, illustrated in Kiew (2000). 

Sotes: This is one of the most decorative of Sabah begonias with its polished 
red stems that give them a lacquered appearance, the dainty pointed leaves 
with a crimson scalloped margin and the many carmen, heart-shaped male 
buds set on slender sprays. It would be a fine addition to the cultivated 
begonias and. considering the harsh conditions under which it grows, would 
probably mean it is easier to cultivate than the forest species that grow in 
deep shade. 

Specimens examined: SABAH: Semporna District - Batu Tengar Cave (Segarong Cave) 
Symington & Agama 9312 (K. SING). Keith A9416 (K. SING). ~Kiew et al. RK 4327 (SAN. 
SING). 

LL Begonia lambii Kiev*, sp. now 

Section Petermannia 

A Begonia inostegia Stapf foliis dense hirsutis obovatis brevipetiolatis et 
fructibus oblongis differt — TYPUS: Batu Tinahas R. Kiew & S. 
Anthonysamy RK 4405 (holo SAN: iso K. SING). 



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Card. Bull. Singapore 53 (2001) 




Figure 5. Begonia keithii Kiew 

A Habit x '/ 2 , B & C Branchlet of male inflorescence x V 2 and x 1, D Male flower bud x 1 V : , E 
Male flower x 1V 2 , F. Androecium x 5, G Stamen x 10, H Female flower x 1, 1 Open female 
flower x l'A, J Style and stigma x 5, K T.S. ovary x 2, L Fruit x 1. (from RK 4327) 



Limestone Begonias o f Sabah. Borneo 



269 



Figure 6 

Rosette herb with 3 — 4 leaves held ± flat on the leaf litter layer. Stem and 
petioles with brown and scurf)* indumentum giving a felted appearance 
with additional sparse long hairs. Stem unbranched. becoming woody, erect, 
to 10 — 13 cm tall and 3 — 5 mm diam.. internodes 1.25 — 2.5 cm long. Stipules 
pale green, lanceolate, c. 14 x 14 mm. margin entire, midrib keeled, apex 
cuspidate, caducous. Leaves alternate, distant, held horizontally: petiole 
1 — 2.5 cm. shallowly grooved above: lamina plain dark green above, green 
or reddish brown or maroon beneath, hairs on upper surface upstanding, 
eglandular. uniseriate. c. 1.5 mm long, translucent in completely green 
leaves, reddish brown, purple or magenta where lamina is coloured beneath, 
glabrous beneath, thin. matt, broadly obovate. strongly asymmetric. 12 — 
14.5 cm x 7.5 — 11cm, narrower side slightly concave, broader side 5 — 7.5 
cm wide rounded, base unequally cordate, basal lobe 1 — 3 cm long, margin 
denticulate with each tooth tipped by long hair, apex rounded or shortly 
acuminate (acumen -I — 5 mm long): venation palmate, midrib and veins 
brown or deep purple and densely hairy beneath. 5 — 7 veins, midrib ± 
same size as the laterals, branching dichotomously 3 — 4 times before 
reaching margin, c. 3 veins in larger basal lobe, veins deeply impressed 
above giving the lamina a corrugated appearance. 

Inflorescence protogynous. from axil of new leaf, densely hairy, erect, longer 
than petioles, at first compact and cymose 1 — 3 cm long with one female 
flower, then producing a spike 7.5 — 10 cm long of distant cymules with 
crowded male flowers, peduncle 1.25 — 2.5 cm: bracts pale green, narrowly 
lanceolate. 5 x 1.5 mm. entire, persistent. Male flower with white pedicel 
A — 5 mm long, densely hairy: tepals 2. pure white, densely hairy on outer 
convex central portion, broadly elliptic. 5 — 9 x 4.5 — 8. entire, apex rounded: 
stamens c. 30. cluster conical. 2 — 3.5 x 3 — 3.5 mm. usually sessile, filaments 
0.5 — 0.75 mm long, anthers obovate. c. 1.5 x 0.75 mm. apex not emarginate. 
Female flower: pedicel c. 4 — 7 mm long, white, densely hairy: ovary green, 
sparsely hairy on the wings, densely hairy on the locules. oblong with 
wings expanding slightly distally. 5 — 9 x 3 — 8 mm. wings 3. reddish, 
isomorphic, wings c. 1 — 2 mm wide, locules 3. placentas bilamellate: tepals 
5. white, outer surface with long hairs. ± isomorphic, innermost slightly 
narrower, elliptic. 9 — 12 x 5 — 8 mm in outer 4 tepals and 4 — 5 mm wide in 
innermost tepal. entire, apex acute: styles 3. greenish, c. 4 mm long, joined 
for c. 1 mm at base, distally bifurcating, stigma papillose forming a 
continuous twisted band. Fruit decurved and pendent, pedicel rigid. 1 — 2 
cm long, capsule oblong, rounded at base, truncate or rounded at apex. 
15 — 18( — 25) x 10 — 13( — 14) mm. hispid particularly on locules. style 
caducous, locules 3. dehiscing between locule and wing, wings 3. equal. 



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Gard. Bull Singapore 53 (2001) 



thin, 2 — 4 mm wide; seeds brown, minute, broadly ellipsoid, c. 0.2 — 0.25 
mm long, tapered to base, rounded distally. 

Distribution: Borneo - SABAH: Pensiangan District, Batu Punggul and 
Batu Tinahas; Sapulut. 

Habitat: Steep earth slopes up to or on flat areas at the base of limestone 
cliffs. 

Notes: The species is striking in several respects. Its deeply corrugate, 
obovate leaves with a dense covering of magenta or purple hairs contrasting 
with the dark green lamina make it a distinctive species. Its habit too, 
where the dark green leaves lie on and blend with the leaf litter, is also 
unusual. It is illustrated in Kiew (1998d, 2000). 

It is a typical member of Section Petermannia in having an upright 
habit, being protogynous, the female flowers having 5 sepals, a 3-loculate 
ovary with bilamellate placentas, the male flowers having 2 tepals, obovate 
anthers that are not emarginate, and the capsule having 3 isomorphic wings 
and caducous styles. 

In inflorescence structure, it is closely similar to B. inostegia Stapf 
from Gunung Kinabalu. Both at first have a short inflorescence, which 
produces the single female flower, as this matures into the fruit, an erect 
axis is produced with a few distant, short lateral branches that produce 
bunches of crowded male flowers. The inflorescence is thus protogynous 
and the single female flower is basal to the many male flowers. However, 
the leaves of these two species are very different, those of B. inostegia are 
long-petioled, ovate with a cuspidate apex, and the capsule is obovoid. In 
addition, B. lambii also does not have the peculiar ovate, fimbriate stipules 
characteristic of B. inostegia. 

This decorative species is named for Anthony Lamb of the 
Agricultural Park, Sabah, whose deep interest in Sabah plants has inspired 
him to bring this species (and many others of horticultural potential) into 
cultivation. 

Specimens examined: SABAH: Pensiangan District - Batu Punggul Ali Ibrahim A I 135 
(SING). Sumbing Jimpin SAN 135991 (SAN), Batu Tinahas the type and R. Kiew & S. 
Anthonysamy RK 4408 (K, SAN. SING); Sapulut (near Kampung Naaturan) R. Kiew & S. 
Anthonysamy RK 4345 (SING). It is also cultivated at the Agricultural Park. Tenom. 



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271 




Figure 6. Begonia lambii Kiew 

A Habit x 'A, B Male inflorescence x 1 , C Male bud x 7 4 , D Open male flower x 1, E Androecium 
x 4. F Stamen x 10, G Female flower x 1, H Open female flower x 1,1 Style and stigma x 4, J 
T.S. ovary x 1V 2 , K Fruit x 1. (from RK 4408) 



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Gard. Bull. Singapore 53 (2001) 



12. Begonia layang-layang Kiew, sp. nov. 

Section Petermannia 

A Begonia amphioxo Sands foliis latioribus et fructibus majoribus pedicello 
longiore suffultis — TYPUS: Sapulut R. Kiew & S. Anthonysamy RK 4441 
(holo SAN; iso BRUN, K, L, SAR, SING) 

Cane-like, glabrous begonia. Stem pale green at first, becoming brown, c. 1 
m tall and 3 — 6 mm diam., woody, internodes up to 4 cm long, erect, 
branching dichotomously, young branches conspicuously zigzag, nodes 
annular. Stipules pale green, narrowly lanceolate, to 12 — 14 x 2 — 4 mm 
with prominent midrib, margin entire, apex acute, caducous. Leaves 
alternate, distant, pendent; petiole pale green, (2.5 — )3.5( — 5.5) cm and in 
dried state 1 mm thick, slightly grooved above; lamina plain dark green 
above except for the red patch at point of attachment with petiole, pale 
green or sometimes reddish beneath, succulent drying slightly leathery, 
slightly glossy above, peltate, slightly asymmetric, narrowly elliptic, (8 — ) 
10(— 11.5) x (2— )2.2(-^.5) cm, the broader side (1.7— )2(— 2.7) cm wide, 
basal lobe acute to pointed sometimes slightly oblique, length of basal lobe 
(3.5 — )4.2(— 5.5) cm, margin red with fine distant teeth becoming closer 
and more prominent towards apex, apex attenuated; venation palmate- 
pinnate, midrib and lateral veins concolorous with lamina, 1 — 4 unbranched 
pairs, 3 veins in basal lobe, plane above, slightly prominent beneath. 
Plant protogynous. Male inflorescences from upper leaf axils, green, erect, 
longer than petioles, a slender, lax three times-branched panicle with c. 15 
flowers, 3.7 — 5.2 cm long of which peduncle is 2.2 — 3.7 cm; bracts lanceolate, 
thin, green, lower ones c. 12 x 3 mm, becoming smaller towards stem apex, 
midrib prominent, margin entire, caducous. Male flowers with whitish pedicel 
(3 — )5 — 7 mm long, tepals 4, outer two pinkish outside, white inside, 
glabrous, subrotund, c. 6 x 5.5 mm, entire, apex rounded, inner two white, 
narrowly elliptic, c. 5 x 1.75 mm, apex rounded; stamens over 50, cluster 
conical, sessile, c. 3 x 2.5 mm, filaments c. 1 — 1.25 mm, anthers golden 
yellow, obovate, c. 0.75 — 1 mm long, apex emarginate. Female flowers 
solitary (rarely 2 per axil) in lower leaf axils; pedicel 17 — 27 mm long; 
ovary deltoid. 17 — 27 x 19 — 30 mm, wings 3 equal reddish, c. 4 — 8 mm 
wide, locules 3, placentas axillary, bilamellate with many ovules on both 
surfaces; tepals 5, free, greenish white, outer four broadly elliptic, slightly 
tapered to base, 9 — 15 x 5 — 9 mm, entire, apex rounded, inner one narrower, 
slightly obovate, 10 — 12 x 4 — 5 mm, apex rounded; styles 3, dark yellow, 
4 — 5 mm joined for c. 1.75 mm at base, distally bifurcating; stigma papillose 
forming a continuous twisted band. Fruit dangling, pedicel extremely fine, 
(1.3 — )2.8( — 3.7) cm, capsule broadly deltoid, distal edge straight, (20 — ) 



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273 



24.5( — 29) x (22 — )26( — 33) mm. locules 3, dehiscing between upper half 
of wing and locule. wings 3. isomorphic, rounded proximally, truncate 
distally. 7( — 8) mm wide, thinly fibrous, style caducous; seeds brown, minute, 
broadly ellipsoid, c. 0.3 mm long, base narrowed, rounded distally. 

Distribution: Borneo - SABAH: Pensiangan District. Sapulut. 

Habitat: In shade below tree canopy on rocks at base of cliff and on 
subsummits. 

Notes: It is a typical member of sect. Petermannia having a cane-like habit, 
being protogynous and having fruits with three equal wings and a caducous 
style. 

In its peltate and relatively narrow leaves, it most resembles B. 
amphioxus. However, it is clearly distinct from this species in its non- 
variegated leaves (those of B. amphioxus are red spotted), which are broader 
and less than four times longer than wide (compared with 8 — 28 mm wide 
and at least five times longer than wide in B. amphioxus), the tepals of the 
female flower are always free (frequently joined in B. amphioxus). and the 
much larger fruits on long pedicels with three broad equal wings. The 
fruits of B. amphioxus are smaller. 9 — 13 x 5 — 14 mm. frequently are two- 
winged, the wings are only 2 — 5 mm wide and the pedicels 2 — 2.5 cm long. 
(Sands, pers. comm., informs me that 2 — 2.5 mm given as pedicel length in 
his type description is in error and should read 2 — 2.5 cm). 

Both B. layang-layang and B. amphioxus grow in moderate shade on 
rock faces quite close to the base of the cliff. In this they differ from B. 
baturongensis. B. keithii and B. madaiensis that grow fully exposed or in 
light shade and do not flower in deep shade. 

This species takes its name from the local one, layang-layang meaning 
swallow in Malay, because of the resemblance of the leaf shape to a 
swallow's wings. It is illustrated in Kiew (1998d) as "the swallow begonia'. 

Specimens examined: SABAH: Pensiangan District - Sapulut (near Kampung Labang) the 
type and Diwol Sundaling SAN 13522 (SAN). Aban & Singh SAN 107961 (SAN). 

13. Begonia madaiensis Kiew, sp. nov. 

Section Petermannia 

A Begonia baturongensi Kiew fructibus longioribus quam latioribus apice 
acutis — TYPUS: Gunung Madai R. Kiew et al. RK 5057 (holo SAN; iso 
K. SAR. SING). 



274 



Card. Bull. Singapore 53 12001) 



Figure 7 

Cane-like, glabrous begonia with several stems arising from prostrate 
rhizome. Stem reddish brown when young, at first erect, then becoming 
prostrate or pendent if growing on cliffs, to 1.7 m tall x 4 — 10 mm diam., 
internodes 3.5 — 10 cm long, woody, slightly thicker at nodes much branched, 
twigs zigzag to 60 cm long, maroon. Stipules pale rosy purple, narrowly 
lanceolate, c. 20 — 21 x 5 — 6 mm, margin entire, midrib keeled, apex 
attenuate, caducous. Leaves alternate, distant, pendent; petiole maroon, 
1.2 — 5 cm x 1 — 3 mm long, terete; lamina green, dark green or brown- 
green, beneath maroon or pinkish light green, margin red, veins red above 
and beneath, young leaves variegated above with oval whitish-green or 
silvery spots of varying size between the veins, sometimes coalescing, and 
with tiny spots along the margin, veins deep ruby red above and rosy 
purple beneath, expanding leaf strongly pleated, succulent and brittle in 
live state, membranous when dried, surface slightly scintillating and 
appearing finely velvety, obliquely ovate, asymmetrical. (5 — )10( — 13.5) x 
(1.7 — )4( — 6) cm, the broader side being (1 — )2.2( — 3.2) cm wide, basal 
lobe broadly rounded (1.2 — )2.5( — 5) cm long, margin scalloped, apex 
acuminate; venation palmate-pinnate, midrib and lateral veins red beneath, 
(2 — )3 veins branching c. midway to the margin with (1 — )2 veins in the 
basal lobe, plane above, slightly prominent beneath. 
Plant protogynous. Male inflorescences about 5 produced from upper axils, 
brownish red, erect, slightly longer than petioles, once-branched cyme with 
up to 7 flowers, 3.7 — 4 cm long of which 2 — 2.7 cm is the peduncle; bracts 
narrowly lanceolate, c. 12 — 14 x 3 mm, caducous. Male flower with pink 
pedicel 7 — 9 mm; tepals 4, outer two white or rosy pink, glabrous, subrotund, 
(5 — )7(12) x (6 — )7( — 9) mm, margin entire, apex rounded, inner two pure 
white, narrowly obovate, 6 — 7 x 2 mm, margin entire, apex rounded; stamens 
c. 40, cluster broadly ovoid, sessile, c. 4 x 3 mm, filaments 0.5 — 0.75 mm, 
anthers golden or pale yellow, obovate, c. 1 mm long, apex emarginate. 
Female flowers solitary from lower leaf axils, up to 5 produced; pedicel 9 — 
11 mm long, pale red; ovary pale green-white with edge of wings reddish, 
oblong, c. 11 — 12 x 9 mm. wings 3, equal, locules 3, placentas axile, 
bilamellate with many ovules on both surfaces; tepals 5, white faintly pink 
towards edge, elliptic, margin entire, apex acute, isomorphic, the inner 
slightly narrower, outer four c. 9 x 6 mm, inner one c. 9 x 4 mm; styles 3. 
yellowish green or golden yellow, 4 mm long joined for c. 0.5 mm at base, 
bifurcating: stigmas papillose forming a continuous twisted band. Fruit 
pedicel withering and becoming thread-like so the fruit dangles, (6 — )14( — 
20) mm long, capsule obovate, (17— )25(— 29) x (10— )18(— 20) mm, 
glabrous, locules 3. dehiscing between locule and wing in the upper half of 




Figure 7. Begonia madaiensis Kiev 

A Habit x 'A. B Stipules x 1. C Branchlet of male inflorescence x 2, D Androecium x 5. E 
Stamen x 15, F Fruit x 1. (from LSP 672) 



276 



Card. Bull. Singapore 53 (2111)1 ) 



capsule, wings 3, equal, rhomboid, (3 — )5( — 7) mm wide, thinly fibrous; 
seeds brown, minute, broadly ellipsoid, c. 0.25 — 0.3 mm long, base truncate, 
distally rounded. 

Distribution: Borneo - SABAH: Lahad Datu District, Gunung Madai. 

Habitat: Growing directly on limestone rocks and cliff faces in light shade. 

Notes: In its erect habit, being protogynous with solitary female flowers 
below and male inflorescences above, in the 3-loculate ovary with 
bilamellate placentas and capsule with 3 equal wings and a caducous style, 
it is typical of sect. Petermannia. However, the male flowers have four 
tepals. 

Among the cane-like begonias on limestone in Sabah, B. madaiensis 
is most similar to B. baturongensis in their leaves being broader and having 
a rounded basal lobe compared with those of B. amphioxus and B. keithii, 
and in being smaller than those of B. heliostrophe, B. keeana and B. 
malachostica. B. madaiensis can easily be told apart from B. baturongensis 
by its fruit, which narrows to the apex, giving it a rhomboid outline. Besides 
fruit shape, it also differs from B. baturongensis in several other features, 
such as its scalloped leaf margin, smaller bracts, less branched male 
inflorescence, white or pale pink flowers, tepals of the male flower that are 
longer than broad, and the leaves, which are strongly pleated before they 
expand. 

The leaves of young plants are larger (about twice the size of those 
on fertile shoots) and can be conspicuously variegated, but this coloration 
is not expressed in older leaves. Plants growing in deeper shade tend to 
have greener, less maroon leaves and plants in deep shade, i.e. at the base 
of cliffs, do not flower. Flowering on a single twig is effectively unisexual 
as the female phase does not overlap with the later male phase. 

Specimens examined: SABAH: Lahad Datu District - Gunung Madai the type and W. 
Meijer SAN 37950 (SAN), SAN 37959 (SAN). S.P. Lim el al. LSP 672 (SAN. SING), LSP 
702 (SAN, SING). 

14. Begonia malachosticta Sands 

Section Petermannia 

Sands, Kew Magazine. 7 (1990) 64 & Plate 145. 

TYPE: Bukit Dulong Lambu Sands & Young - Sands 3933 (not seen) 

Distribution: Borneo - SABAH: Kinabatangan District, Bukit Dulong 
Lambu (Gomantong Cave). 



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277 



Habitat: In rocky crevices in the sheer cliffs where the tree canopy begins 
to open up, as well as on exposed jagged outcrops. It is no longer common 
on the summit, which is greatly disturbed by the village of birdnest collectors 
that is established on the summit plateau. 

Notes: This is a strikingly beautiful species with pink spotted leaves with a 
red underside. It has been introduced into cultivation at Kew where it is 
grown in a 50:50 mix of peat and sand. It is propagated easily from nodal 
cuttings (Sands. 1990). 

It is eaten as a vegetable but is reportedly very sour [Kamideh SAN 
66578). 

Specimens examined: SABAH: Kinabatangan District - Bukit Dulang Lambu (Gomantong 
Cave) James Ah Wing SAN 38108 (SAN), SAN 47263 (SAN): Kamideh Bangillan SAN 
66578 (SAN). 

15. Begonia melikopia Kiew, sp. nov. 

Section Petermannia 

A Begonia erythrogyna Sands foliis latioribus. inflorescentiis minoribus. 
tepalis rubris et fructibus angustioribus differt — TYPUS: Gua Melikop 
Ruth Kiew et al RK 5011 (holo SAN; iso BRUN, K. L. SAR. SING). 

Cane-like, glabrous begonia with prostrate rhizome with several spaced 
erect stems to 1.5 m tall x 5 — 7 mm diam.. base becoming woody, young 
stems brittle at swollen nodes, reddish above nodes, otherwise green, leaf 
scars very conspicuous, internodes 2.5 — 5.5 cm long, slightly ribbed, stems 
with 1 — 2 branches. Stipules green, strongly keeled, lanceolate, 23 — 25 x 
9 — 11 mm. margin entire, apex acute, caducous. Leaves alternate, distant, 
in a bunch at top of stem, held horizontally: petiole completely reddish or 
reddish proximally and distally but green in between. 8.5 — 13 cm x 4 — 6 
mm, grooved above: lamina plain mid-green above, beneath completely 
magenta or with magenta blotches and green veins, seedling leaves with 
white spots, succulent and brittle in life, thinly chartaceous in dried state, 
glossy above, asymmetric, broadly ovate, sometimes almost reniform, 15 — 
19.5 x 12 — 18.5 cm of which broader side is 7.5 — 11.5 cm wide, basal lobes 
broadly rounded. 3.5 — 7.5 cm long, base cordate but not overlapping, margin 
scalloped between veins and minutely and distantly dentate, apex cuspidate: 
venation palmate-pinnate, midrib and lateral veins concolorous with lamina 
above. 5 — 7 pairs, branching twice dichotomously. first less than halfway 
to margin, then c. halfway to margin. 1 — 2 veins in basal lobe, impressed 
above, prominent beneath. 



278 



Gard. Hull Singapore 53(2001) 



Inflorescence axillary, protogynous with 2 female flowers at base of the 
erect cymosely branched panicle of male flowers, shorter than petioles, 
6 — 7.5 cm long of which peduncle is 1.7 — 2.5 cm with diam. c. 2 mm, 
branching c. 3 — 4 times, c. 4 — 8 main branches 4 — 15 mm long, ultimate 
branches 8 — 20 mm long, knobbly from congested scars of fallen flowers, 
each terminating in 1 bud or 1 bud and 1 one flower. Male flower with 
slender pedicel 7 — 12 mm long, tepals 2, flame-coloured darker towards 
base, glabrous, oval, 8 — 11 x 4 — 5 mm, margin entire, apex rounded; stamens 
c. 25, cluster conical, 3.5 — 4 x 2.5 — 3 mm, sessile, filaments c. 0.75 mm long, 
anthers pale yellow, spathulate, c. 1.2 mm long, apex not emarginate. Female 
flower not known. Fruit pendent and dangling on fine and thread-like 
pedicel 25 — 38 mm long, capsule narrowly deltoid narrowed into pedicel. 
40 — 55 x 10 — 27 mm, glabrous, locules 3, placentas axile, bilamellate with 
many seeds on both surfaces, dehiscing between wing and locule, wings 3, 
equal, 6 — 7 mm wide, becoming thin and papery, tip acute or sometimes 
rounded; seeds brown, minute, almost globose, c. 0.2 mm long, base tapered, 
rounded distally. 

Distribution: Borneo: SABAH - Kinabatangan District, Gua Melikop. 

Habitat: Gua Melikop is an exposed cliff face on a hillside at about 400 m 
altitude. Its summit is covered by a deep layer of peat, which does not 
support a limestone flora. 

Begonia melikopia grows rooted in cracks in the sheer limestone 
rock face in deep shade below the canopy but c. 1.5 — 2 m above base of 
limestone cliff. (Some seedlings were found growing in soil near the base 
of the cliff but no adult plants were found on soil, suggesting that these 
seedlings might not survive to maturity). 

Notes: In its large, broadly ovate leaves, paniculate male inflorescence and 
large deltoid fruits pendent on long pedicels, it resembles B. erythrogyna 
Sands. However, it is distinct from this species in its much broader leaves 
(those of B. erythrogyna are less than 16 cm wide), its shorter, axillary male 
inflorescence (terminal and 7 — 14 cm long in B. erythrogyna), the male 
flowers are flame-coloured (they are white to pale green in B. erythrogyna), 
and the fruit, which is more than twice as long as wide (in B. erythrogyna it 
is less than twice as long as wide). 

Begonia melikopia is typical of sect. Petermannia in its cane-like 
habit, in being protogynous, having male flowers with two tepals, and the 
fruits with three equal wings and three locules each with bilamellate 
placentas. 



Limestone Begonias ofSabah, Borneo 



279 



Specimens examined: Known only from the type population. 

16. Begonia postarii Kiew 

Section Petermannia 

Kiew. Gardens' Bulletin Singapore. 50 (1998) 165 & Fig. 1. 
TYPE: Bukit Panggi Kiew & Lim RK 4221 (holo SAN; iso SING). 

Figure 8 

Distribution: Borneo - SABAH: Kinabatangan District, Kinabatangan 
Valley (Bukit Panggi. Bukit Dulong Lambu (Gomantong Cave) and Subuk 
Estate). 

Habitat: At base of limestone hills (but not on cliff faces) growing in deep 
shade in sheltered damp habitats on soil, low limestone boulders or the 
base of tree trunks. 

Notes: This softly hairy begonia with hairy tepals in the male flower and 
the hairy fruit is quite unlike any other limestone species in Sabah. 

Specimens examined: SABAH: Kinabatangan District - Bukit Dulong Lambu Joseph B. et 
al. SAN 122763 (SAN). Lim S.P. & Ubaldus LSP 802 (SAN, SING). Ruth Kiew & Lim S.P. 
BDL 4 (K, SAN. SAR. SING): Bk. Panggi Ruth Kiew & Lim S.P. RK 4221 (SAN, SING): 
Subak Estate (Lower Kinabatangan River) J. Dransfield et al. JD 5770 (SAN). 

17. Begonia punbatuensis Kiew, sp. now 

Section Petermannia 

A Begonia burbidgei Stapf fructibus minoribus. pedicellis filiformibus 2.5 — 
3 cm longis differt — TYPUS: Pun Batu R. Kiew & A. Berhaman RK 4260 
(holo SAN: iso BRUN. K. L. SAR. SING). 

Figure 9 

Cane-like begonia. Stem robust, dark brown, up to 1.3 m tall and 6 mm 
diam.. erect, little branched, woody, internodes up to 4 — 5.5 cm long, nodes 
conspicuously swollen with conspicuous leaf scars, eglandular uniseriate 
hairs c. 0.5 — 0.75 mm long, scattered on the young stem, becoming 
glabrescent. Stipules pale green, glabrous except for a few scattered hairs 
on the outer surface of the midrib, slightly obovate, 2 — 3.5 x 1 — 1.3 cm, 
margin entire, apex acute, persistent. Leaves alternate, distant: petiole green 
with brown bristly hairs. 2.5 — 6 cm long, grooved above: lamina plain mid- 
green above and beneath except for crimson patch on upper surface at the 
junction with the petiole, succulent and brittle in life, thinly leathery in 



280 



Card. Bull. Singapore 53 (2001) 




Figure 8. Begonia postarii Kiew 

A Habit x '/,. B Branchlet of male inflorescence x 1 'A. C Male bud x 1 . D Male flower x 2, E 
Androecium x 4V 2 , F Stamen x 8. G Fruit x 1. (from RK 3838) 



Limestone Begonias ofSabah, Borneo 



281 



dried state, glossy above, obliquely ovate, strongly asymmetric, 5 — 9 x 6 — 
10 cm, narrow side lanceolate, broad side broadly ovate 4.5 — 7 cm wide, 
base cordate, not overlapping, basal lobe 1.7 — 4 cm long, margin shallowly 
dentate, faintly red in young leaves, apex acute; venation palmate, midrib 
and lateral veins concolorous with lamina, sparsely hairy beneath. 3 — 4, ± 
equal-sized, branching dichotomously twice before reaching the margin, 
first close to petiole, second towards the margin, 1 vein in basal lobe, plane 
above, prominent beneath. 

In florescences protogynous. in the axils of upper leaves, pale green, glabrous, 
± erect, longer than adjacent petiole, cymose, with up to 4 inflorescences 
at successive leaf axils, each branched at base, one axis with 2 female 
flowers on peduncle 2.5 — 3 cm long, the other axis bears the male 
inflorescence with c. 4 lateral branches, conspicuously jointed, 5.5 — 9 cm 
long of which peduncle 3.2 — 4.5 cm long; bracts pale green, obovate, those 
at base c. 20 x 15 mm. those subtending male flowers c. 10 x 9 mm, persistent. 
Male flowers with whitish green, glabrous pedicel 17 mm long; tepals 4, 
rosy or pale pink, glabrous, outer two broadly oval, 14 x 13 or 15 x 10 mm, 
inner two narrowly elliptic, c. 9 x 5 mm, margin entire, apex rounded; 
stamens c. 50, cluster conical, sessile, 9x4 mm, anthers subsessile, golden 
yellow, narrowly oblong, c. 2 x 1 mm, apex emarginate. Female flower with 
pale green, glabrous pedicel; ovary pale greenish white, glabrous, ovoid, 15 
x 15 — 17 mm, wings 3, isomorphic, 3 locules, placentas axile and bilamellate 
with many ovules on both surfaces: tepals 5. pale rosy pink, broadly elliptic, 
outer 17 x 10 mm, inner 14x7 mm margin entire, apex slightly acute; styles 
3, golden yellow, c. 5 — 6 mm long, bifurcating: stigma papillose forming a 
continuous twisted band. Fruits pendent, dangling on fine, thread-like 
pedicel 2.5 — 3 cm long, capsule 17 — 22 x (12— )20 — 28 mm. glabrous, 
locules 3. dehiscing between upper half of locule and wing, wings 3, 
isomorphic, truncate distally (4 — )6 — 7 mm wide, thinly fibrous; seed brown, 
minute, ellipsoid, c. 0.5 mm long, base truncate, distally rounded. 

Distribution: Borneo - SABAH: Keningau District, Pun Batu. 

Habitat: Summit of the limestone hill growing on exposed peat-covered 
rocks. 

Notes: Pun Batu is an isolated tower karst hill and its summit is covered in 
a deep peat layer. Its unique begonia grows in this habitat and as yet has 
not been found on any other limestone hill. The surrounding forests are 
actively being logged but when we visited in 1996, its surrounding foothills 
were still pristine. 

In its upright habit, in being protogynous, and with the fruit being 3- 



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Figure 9. Begonia punbatuensis Kiew 

A Habit x 7 : , B Branchlet male inflorescence x 1. C Male bud x l'A. D Male flower x 1. E 
Androecium x 3. F Stamen x 10, G Fruit x 1 . (from RK 3898 ) 



Limestone Begonias ofSabah. Borneo 



283 



loculate with three equal wings and a bilamellate placenta and caducous 
style, it falls within sect. Petermannia. 

Begonia punbatuensis resembles B. bnrbidgei from Gunung Kinabalu 
in its robust, woody stems with swollen nodes, large stipules on the young 
stems and in its leaf shape and venation. However, B. bnrbidgei is a larger 
plant growing to c. 2 — 2.7 m tall, its leaf apex is acuminate to cuspidate, its 
inflorescences have larger bracts, the male flowers have much larger tepals 
(the outer two tepals measuring c. 25 x 20 mm), and the fruits are also 
much larger (up to 27 x 46 mm), the wings are not truncate and the fruit 
stalk is stiff and short (c. 15 mm long). 

Although its bracts are quite conspicuous. B. punbatuensis does not 
belong to sect. Bracteibegonia sensu Doorenbos et al. (1998), which includes 
just three species from Java and Sumatra. These differ from species in sect. 
Petermannia in having pinnate venation, few flowered bisexual 
inflorescences and persistent styles in the fruit. Doorenbos et al. therefore 
transferred the Bornean species with conspicuous bracts, previously placed 
in sect. Bracteibegonia (such as B. burbidgei Stapf and B. inibricata Sands), 
to sect. Petermannia. to which they conform in their venation, inflorescence 
structure and caducous style. 

Specimens examined: SABAH: Pensiangan District - Pun Batu the type and Ruth Kiew & 
A. Berhaman RK 4279 (E. KEP. SAN. SING). 

18. Begonia urunensis Kiew, sp. nov. 

Section Petermannia 

A Begonia anthonyi Kiew and B. berhamanii Kiew caulibus ramosis, foliis 
latioribus et inflorescentiis masculis paniculatis differt — TYPUS: Batu 
Urun R. Kiew & S. Anthonysamy RK 4473 (SAN, SING). 

Low, erect, bushy herb, trichomes uniseriate c. 0.5 mm long, dense on 
young stems, stipules, petioles, lower surface of veins, inflorescence and 
pedicels, sparse on lamina. Stem to 33 cm tall and 5 — 7 mm diam., woody 
towards the base, lowermost internodes 4.5 — 10 cm long, lower nodes 
swollen, upper internodes 3 — 4.5 cm long, producing short branches which 
sometimes branch again. Stipules lanceolate, 5.5 — 17 x 2.5 — 3.5 mm, margin 
entire, apex attenuate to setose, caducous. Leaves alternate, distant, towards 
apex some leaves subopposite; petiole 6 — 15 mm long in lower leaves, 4 — 7 
mm in the upper; lamina plain mid-green above, (12.5 — )15 — (15.5) x (5 — ) 
6.5( — 8) cm, slightly succulent in life, papery when dry, obovate, sometimes 
rhomboid and widest midway, slightly asymmetric, narrow side concave 
towards the base, broad side rhomboid, 3 — 4.2 cm wide, base unequally 



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cordate, basal lobe 2.5 — 7.5 mm long and auriculate, margin minutely and 
distantly dentate, apex narrowly attenuate; venation pinnate, lateral veins 
6 — 7 pairs, branching once, sometimes twice before reaching the margin, 1 
vein in basal lobe, prominent above and beneath. 

Plant protogynous with 2 female flowers and a male inflorescence from 
each of upper leaf axils. Male inflorescence erect, 0.8 — 2.5 cm long of 
which peduncle is to 1 cm long longer than petioles, central axis with 
several tiers of three times branched side branches c. 7 mm long; bracts 
lanceolate, c. 7 x 2 mm, margin entire, apex setose, caducous. Male flowers 
with pedicel 3 — 5 mm long; tepals 2, pinkish, densely hispid outside, elliptic, 
c. 5 x 3.5 mm, margin entire, apex rounded; stamens c. 40 — 50 in conical, 
sessile cluster; filaments c. 0.5 mm long; anthers orange, spathulate, c. 0.75 
mm long, apex not emarginate. Female flower not known. Fruit with 
decurved, stiff pedicel (4 — )7( — 9) mm long; capsule broadly oblong, (14 — ) 
18( — 21) x (15 — )17( — 19) mm, glabrous, locules 3, placentas axile, 
bilamellate with many seeds on both surfaces, dehiscing along entire length 
between locule and wing, wings 3, isomorphic, tip rounded, (4 — )6( — 7) 
mm wide, thinly fibrous; seed brown, minute, broadly ellipsoid, c. 0.2 mm 
long, base truncate distally rounded. 

Distribution: Borneo - SABAH: Pensiangan District, Batu Urun. 

Habitat: Deeply shaded forest on soil on steep slopes up to and at the base 
of limestone. 

Notes: Its erect habit, three locular ovary and fruit with three equal wings 
place this species within sect. Petermannia. Among the limestone species, 
it groups with B. anthonyi and B. berhamanii, which are short begonias 
with bristly stems, their leaves are only slightly asymmetric, the petioles 
short and the midrib more or less in line with the petiole, the male flowers 
have two tepals, and the fruits do not have long, thin, thread-like pedicels. 
However, it is not typical of this group of begonias in two respects. Firstly, 
it differs in habit. In B. urunensis, the stem branches once or sometimes 
twice to produce flat sprays of leaves giving it a bushy appearance. (The 
stems of B. anthonyi and B. berhamanii are unbranched). Secondly, the 
male inflorescences are produced from the same leaf axil as the female 
flowers and are erect and paniculate, whereas in B. anthonyi and B. 
berhamanii, the female flowers are produced from the lower axils and the 
short, spicate male inflorescence from the upper. B. urunensis can also be 
told apart from these two species by its broader leaf, which is less than 2.5 
times longer than broad as compared with 2.7 — 4 and 5 times, respectively 
for B. anthonyi and B. berhamanii. 



Limestone Begonias of Sabah. Borneo 



285 



Batu Urun is one of the most remarkable limestone formations in 
Sabah being unique in its cave, which has its mouth on the side of a deep 
and wide basin below soil level. From the mouth of the cave a dangerouslv 
steep scree slopes precipitously down to a subterranean river, which to 
judge from the large tree trunks lodged in it carries strong currents in rainy 
weather. On one side of the basin is a vertical wet cliff face that is covered 
in an undescribed species of Monophyllaea (Gesneriaceae), one of two 
Monophyllaea species to be found on limestone in Sabah (Kiew.l998a). 
Above soil level surrounding this basin is a wide area where the limestone 
projects just above the soil surface, forming an uneven and pitted rock 
pavement, or it outcrops as low cliffs up to 4 — 5m high or as boulders of 
various sizes, all in deep shade below the canopy. It is on soil over limestone 
that this new begonia species is found. 

Specimens examined: SABAH: Pensiangan District - Batu Urun. the type and Ruth Kiew & 
S. Anthonysamy RK 4472 (K. SAN. SING). 



Acknowledgements 

This paper forms part of the results from "The Biodiversity and Conservation 
of the Limestone Flora of Sabah* project carried out in collaboration with 
the Forest Research Centre (FRC). Sabah Forest Department. The author 
thanks the Singapore Botanic Gardens and the Ministry of Science, 
Technology and Environment. Malaysia (under IRPA Grant No. 52858) 
for sponsorship; the staff at FRC, Robert C. Ong. K.M. Wong. Diwol 
Sundaling. A. Berhaman and S.P. Lim. for their logistic help and enthusiastic 
support; Anthony Lamb. Herbert Lim and Andy Surat of Agricultural 
Park. Tenom. for showing us localities in the Pensiangan District: J.H. Tan 
with whose support Melikop was located; Mdm P.H. Yap and Jaap J. 
Vermeulen for their excellent botanical illustrations; Andrea Kee for 
carrying out horticultural trials; to the Curators of the BM. E and SAN 
herbaria for permission to examine specimens in their care; to Mark J.E. 
Coode for comments on the manuscript and for correcting the Latin and 
to Martin J.S. Sands for his detailed and helpful comments. 

References 

Beaman, R.S., J.H. Beaman, C.W. Marsh & P.V. Woods. 1985. Drought 
and forest fires in Sabah in 1983. Sabah Society Journal. 8: 10 — 30. 



Doorenbos. J.. M.S.M. Sosef and J.J.F.E. de Wilde. 1998. The Sections of 



286 



Card. Bull. Singapore 53 (2001) 



Begonia Including Descriptions, Keys and Species Lists. Wageningen 
Agricultural University Paper 98-2: 1 — 266. 

Kee, A. 2000. Collecting begonias from the wild. Gardenwise. 15: 14 — 15. 

Kiew, R. 1991. The limestone flora. In: R. Kiew (ed.). The State of Nature 
Conservation in Malaysia. Malayan Nature Society, Malaysia, pp. 42 — 
50. 

Kiew, R. 1998a. Towards a limestone flora of Sabah. Paper presented at 
B.C. Stone Memorial Symposium, Kuala Lumpur. Malaysia. 

Kiew, R. 1998b. Niche partitioning in limestone begonias in Sabah, Borneo, 
including two new species. Gardens' Bulletin Singapore. 50: 161 — 169. 

Kiew, R. 1998c. The unique elements of the limestone flora of Batu Tengar 
Cave (Segarong), Sabah, Malaysia. Gardens' Bulletin Singapore. 50: 185 — 
196. 

Kiew, R. 1998d. Wanderings in the great forests of Borneo. Gardenw ise. 
11: 8—11. 

Kiew, R. 2000. Decorative limestone begonias from Sabah. Gardenwise. 
15: 12—13, 15. 

Lim, S.P. & R. Kiew. 1997. Gazetteer of limestone localities in Sabah, 
Borneo. Gardens' Bulletin Singapore. 49: 111—118. 

Reza Azmi and R. Kiew. 1998. Begonia lazat (Begoniaceae), a new culinary 
begonia from Borneo. Gardens' Bulletin Singapore. 50: 43 — 48. 

Sands, M.J.S. 1990. Six new begonias from Sabah. Kew Magazine. 7: 57 — 
85. 

Sands, M.J.S. 1996. Begoniaceae. In: M.J.E. Coode. J. Dransfield, L.L. 
Forman, D.W. Kirkup and Idris M. Said (eds.) A Checklist of the 
Flowering Plants and Gymnosperms of Brunei Darussalam. Forestry 
Department, Brunei & Royal Botanic Gardens. Kew. pp. 38 — 40. 

Tebbit, M.C. 1999. A revised classification of selected Asian begonias based 
on evidence from morphology and molecules. In: S. Andrews, A.C. Leslie 
and C. Alexander (eds.). Taxonomy of Cultivated Plants. Royal Botanic 
Gardens, Kew. pp. 401—402. 

Tebbitt, M.C. and CM. Maciver. 1999. The systematic significance of the 
endothecium in Begoniaceae. Botanical Journal Linnean Society. 131: 
203—221. 



Gardens' Bulletin Singapore 53 (2001) 287-289. 



Begonia conipila Irmsch. ex Kiew (Begoniaceae) from 
the Gunung Mulu National Park, Sarawak, Malaysia 

RUTH KIEW 

Singapore Botanic Gardens, 1 Cluny Road, Singapore 259569 

Abstract 

Irmscher*s name. Begonia conipila. is validated for the striking species that grows on the 
Melinau limestone in Sarawak. Borneo. 

Introduction 

The scientific name of this most distinctive species was in fact never validly 
published although, following Irmscher's annotated specimens, it has 
become the accepted name and specimens of it are correctly identified in 
herbaria and the name has appeared in print (Anderson & Chai, 1982; 
Jangowe, 1999). It is timely therefore to validate the name of his species, 
particularly because it is seen by most visitors who climb Gunung Api in 
the Gunung Mulu National Park. 

Low's specimen is selected as the lectotype as it is the earliest 
collection and was annotated as a new species by Irmscher. 

Begonia conipila Irmsch. ex Kiew 

Section Petermannia 

A Begonia havilandii Ridl. foliis minoribus, tepalis masculis 2 et fructibus 
oblongis differt - LECTOTYPUS: -Mooloo' (Gunung Mulu National Park) 
Low s.n. (K). 

Creeping herb or low root climber, roots fibrous produced at nodes in 
contact with substrate, indumentum of stiff, white, uniseriate, eglandular 
trichomes, c. 2 — 5 mm long, each on a raised conical base, dense on young 
stem, petiole, on stipule margin and outer surface of stipule midrib, on the 
lamina dense on the upper surface, margin and lower surface of veins, 
some on lower lamina surface. Leafy stem 15 — 25 cm long and 2 — 3( — 4) 
mm diam., little branched, internodes 2.5 — 5 cm long. Stipules lanceolate 
with a distinct midrib. 7 — 12 x 4 — 7 mm, margin entire, apex setose, 
persistent. Leaves alternate, distant, petiole 1 — 2 cm in upper leaves 
elongating to 2 — 5 cm in lower leaves, lamina somewhat bullate, variegated, 



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Card. Bull. Singapore 53 (2001) 



upper surface dark green to blackish purple, paler between veins, the lower 
surface reddish, rarely light green, in dried state thin and papery, conical 
hair bases giving a pustulate appearance above, asymmetric, broadly ovate 
to reniform, 4 — 7.5 x 3.5 — 8.5 cm, broader side (2.25 — )3.5 — 5 cm wide, 
base rounded to cordate, lobes slightly overlapping, basal lobes rounded, 
(0.75 — )2 — 3.5 cm long, margin irregularly serrate, each tooth setose, apex 
cuspidate; venation palmate, main veins 3, equal-sized, branching c. halfway 
to margin, with another 1 — 2 veins in basal lobes, impressed above and 
prominent beneath. 

Inflorescence axillary, whitish or light green, hispid, erect, longer than 
petioles, racemose, 3.5 — 9.5 cm long of which peduncle is 1.5 — 6 cm, 
protogynous with a single basal female flower and many male flowers 
distally on 2 short lateral branches c. 3 — 11 mm long and crowded in a 
terminal cluster; bracts similar to stipules, c. 5 — 6 x 3 mm, persistent; 
hracteoles light green or whitish, broadly ovate, 0.75 x 0.25 mm, apex 
setose, persistent. Flowers with white, translucent tepals. Male flower with 
whitish or light green pedicel, c. 1 — 3 mm long; tepals 2, sparsely hispid on 
upper half of outer surface, broadly elliptic, c. 5 x 3.5 — 4.5 mm, margin 
entire, apex rounded; stamens yellow, c. 10 — 11, in hemispherical cluster 
joined in a torus c. 0.5 — 0.75 mm, filaments between 0.25 mm on lower 
stamens to 1 mm long in the distal ones, anthers bright yellow, obovate, c. 
0.75 — 1 mm long, emarginate, opening by pores. Female flower with pedicel 
c. 3 — 5 mm long, light green, finely hispid; ovary narrowly oblong, c. 7.5 — 
11 x 3 — 5 mm, wings 3, subequal, two shorter 1.5 mm and broader one 2 
mm wide, locules 3, placentas axile, bilamellate with many ovules on both 
surfaces; tepals 5, isomorphic, narrowly lanceolate, c. 7 — 7.5 x 3 — 4 mm, 
margin entire, apex acute, sparsely hispid on outer surface; styles 3, free to 
base, 5 mm long, bifurcating; stigma greenish, papillose forming a continuous 
twisted band. Fruit pedicel stiff, decurved, (2 — )4 — 6( — 8) mm long, capsule 
oblong, c. (11 — )18 — 20 x 10 — 15 mm, sparsely hispid, locules 3 but one 
scarcely developed, dehiscing between wing and locule, wings thin, narrowed 
to base, truncate distally, two narrower wings, c. 3 — 5 mm wide, broader 
wing c. 5 — 7 mm wide, style not persistent. 

Distribution: Borneo - SARAWAK: endemic to the Melinau limestone in 
the Gunung Mulu National Park (on Gunung Api and low limestone 
outcrops along the Sungai Melinau). 

Habitat: It grows from the base of limestone outcrops to c. 1 170 m in lower 
montane limestone forest on Gunung Api, usually on limestone boulders 
but also on steep screes in deeply shaded habitats. 



Begonia conipila from Sarawak 



289 



Notes: Begonia conipila is typical of section Petermannia in possessing 
upright (although weak) stems (as opposed to prostrate rhizomes), 
protogynous inflorescences with the female flower basal and the male 
flowers distal, male flowers with two tepals and obovate anthers that open 
by pores, and female flowers with five tepals, three free styles and a 3- 
loculate ovary with bilamellate placentas. The fruit is interesting in that 
one locule is hardly developed (it is about a quarter the size of the other 
two). A similar situation is seen in B. amphioxus (Kiew, 2001). 

In its creeping stem with erect branches, broadly obovate leaves with 
stiff pale hairs, serrate leaf margin with each tooth tipped by a long hair, 
the few stamens in the male flowers, and the 3-loculate ovary with 
bilamellate placentas, it resembles B. havilandii Ridl. from Sarawak. 
However, it is distinct from this species as B. conipila has smaller leaves 
(they are c. 6 — 10 cm long in B. havilandii), male flowers with two tepals 
(four in B. havilandii) and an oblong capsule (orbicular in B. havilandii). 

With its small, variegated leaves, it is as Low noted on his specimen, 
'a charming plant'. Jangowe (1999) illustrates this species with a fine colour 
photograph. 

Specimens examined: SARAWAK: Gunung Mulu National Park - Low s.n. 'Mooloo' (K); 
Gunung Api - Abang Moktar S49606 (SAN. SAR). Anderson S4287 (K. SAR). S30732 
(SAR. SING), B.L. Burtt B8354 (SAR). P. Chai S30054 (SAR). S35790 (SARj, C. Hansen 
102 (SAN, SAN), P.J. Martin S38257 (SAR), /. Nielsen 502 (SAR). B.C. Stone 13693 
(SAR). Yii P C. & Abu Talib S58845 (SAR); limestone on Sungai Melinau Chew Wee Lek 
CWL 1055 (SAR, SING), B.C. Stone 13598 (KLU, SAR). 

Acknowledgements 

The author thanks the curators of the herbaria at BM, E, K, KLU, SAN 
and SAR for permission to examine specimens in their care, to Mark J.E. 
Coode for his patience in correcting the Latin, and to M.J.S. Sands for 
comments on the manuscript. 

References 

Anderson, J.A.R. and P.P.K. Chai. 1982. Gunung Mulu National Park. 
Sarawak. 9. Vegetation. Sarawak Museum Journal. 30: 195 — 206. 

Jangowe. J. 1999. Begonias of the national parks of Malaysian Borneo. 
Begonian. July/Aug: 144 — 145. 

Kiew, R. 2001. The limestone begonias of Sabah, Borneo - Flagship species 
for conservation. Gardens' Bulletin Singapore. 53: 241 — 286. 



Gardens Bulletin Singapore 53 (2001) 291-313 . 



New Species, Varieties and Reductions in Diospyros 
(Ebenaceae) in Borneo and Peninsular Malaysia 
including Peninsular Thailand 

FRANCIS S.P. NG 
7o Forest Research Institute Malaysia, Kepong, 52109 Kuala Lumpur, Malaysia 

Abstract 

In the genus Diospyros. seven new species (D. beccarioides Ng, D. brainii Ng, D. crockerensis 
Ng, D. keningauensis Ng. D. lunduensis Ng. D. multinervis Ng and D. parabuxifolia Ng) 
and six new varieties {D. curranii Men. var. kalimantanensis Ng: D. ferruginescens Bakh. 
var. rufotomentosa Ng; D. lanceifolia Roxb. var. iliaspaiei Ng. var. renageorgei Ng, var. 
saliciformis Ng: D. penibukanensis Bakh. var. scalarinervis Ng) are described. Thirty species 
or varieties are reduced to synonymy. 

Introduction 

In revising the genus Diospyros for the Tree Flora of Sabah and Sarawak, 
I took the opportunity to review the genus for Borneo and Peninsular 
Malaysia. This has resulted in the recognition of seven new species and six 
new varieties, and the reduction of 30 species or varieties to synonymy. 

New Species 
1. Diospyros beccarioides Ng, sp. nov. 

Arbor ad 20 m alta; rami dense rubro-brunnee pubescentes demum 
glabrescentes. Folia membranacea ad chartacea, glabra, oblonga ad ovato- 
oblonga 16-30 cm longa 5.5-11 cm lata, basi cuneata leviter attenuata rare 
rotundata, apice acuminato, costa supra immersa plana vel 'marginibus' 
elevatis provisa. infra nervis lateralibus prominentibus paribus 7-13 
incurvatis ante margines anastomosantibus venam intra-marginalem plus 
minusve distinctam formantibus. venatione intercostali prominula laxe 
scalariformi: petiolus 0.8-1.5 cm longus. Inflorescentia mascula cymis 
subsessilis condensatis floribus ut videtur 3 vel plus sed ignotis. Fructus 1- 
3 pedicellis 0.3-0.8 cm longis suffulti, globosi ad 2.5 cm diam. glabri, calyce 
in lobis 4 vel 5 erectis folioso-coriaceis plicatis accrescentibus ad c. 2 cm 
longis 1.5 cm latis diviso. Typus: Yii et al. S 50325, Sarawak (KEP holo. 
SAR iso). 



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Card. Bull. Singapore 53 (2001) 



Figure 1 

Tree to 20 m tall. Twigs densely reddish brown pubescent, becoming 
glabrous with age. Leaves membranaceous to chartaceous, glabrous; oblong 
or oblong-obovate, 16-30 x 5.5-11 cm, base cuneate and slightly attenuate, 
rarely rounded, apex acuminate; midrib above sunken, flat, or sunken with 
raised sides; lateral veins prominent below, 7-13 pairs, inarching and 
anastomosing at the margins to form a more-or-less distinct intramarginal 
vein; intercostal venation prominulous below, laxly scalariform; petiole 8- 
15 mm long. Male inflorescences subsessile condensed cymes of 3 or more 
flowers. Flowers unknown. Fruits 1-3, on 3-8 mm long stalks, globose, to c. 
2.5 cm diam., glabrous. Fruit calyx divided into 4-5 valvate lobes, the lobes 
erect, leafy-coriaceous, plicate, accrescent, to c. 2 x 1.5 cm. 

Distribution and habitat: Sumatra (Jacobs 8341, Lampung Prov.); Borneo, 
Sarawak (S 8633, S 16177, S 31817, S 32625, S 50325, S 65187; Chew CWL 
1100, 1141); Kalimantan (Endert 5164, 5380; Kessler et al. 824, 969); Sulawesi 
(Vogel & Vermuelen 6981). In Sarawak, the species is found on limestone 
hills, up to 200 m altitude. 

Notes: Bakhuizen in Bull. Jard. Bot. Btz. Ser. iii. 15 (1938) 239 had included 
specimens of three species under Diospyros beccarii Hiern. The two 
syntypes of D. beccarii Hiern belong to D. sumatrana Miq., hence D. beccarii 
has to be reduced to D. sumatrana. At least two other of his cited specimens, 
bb. 12051 and bb. 14750, belong to D. daemona Bakh.. The third element, 
represented by, for example, Endert 5164 belongs to this new species, D. 
beccarioides, which can be distinguished from D. sumatrana by its larger 
fruits with accrescent calyx lobes up to 2 cm long, and larger leaves, with a 
stronger tendency to form an intramarginal vein. 

2. Diospyros brainii Ng. sp. nov. 

Arbores mediocres ad 29 m altae. Folia mediocra ad magna, oblongo- 
ovata, 11.5-19.5 x 2.5-6 cm, apice acuminata, basi cuneata, lamina chartacea, 
glabra, costa supra immersa; nervi secundarii infra prominuli ad prominentes 
8-10 pares intus curvati in nervo intramarginali inconspicuo ad circa tertiam 
spatiam inter marginem costamque anastomosantes; nervi tertiarii 
inconspicui reticulati; petiolus 0.5-1 cm. Flores ignoti. Fructus solitarius, 
sessilis, globosus ad 4.5 cm diam., glaber, in sicco valde rugosus. Calyx 
accrescens, in lobis 4 vel 5 coriaceis valvatis 1.5 cm longis profunde divisus, 
in quoque axe principali sursum curvato fructus curvaturam secundo sed 
marginibus inter se deorsum curvatis. Typus: Brain anak Tada, S 15932, 



Diospyros (Ebenaceae) in Borneo and Peninsular Malaysia 




Figure L Diospyros beccarioides Ng. A. leafy twig: and B. fruit. (A-B from S 50325). 



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Card. Bull. Singapore 53 (2001) 




Figure 2. Diospyros brainii Ng. A. leafy twig and B. fruit. (A-B from S 15932). 



Diospyros (Ebenaceae) in Borneo and Peninsular Malaysia 



295 



Borneo. Sarawak (SAR holo. SAN. SING iso). 
Figure 2 

Medium trees to 29 m tall. Leaves medium-sized to large, oblong-obovate. 
11.5-19.5 x 2.5-6 cm. apex acuminate, base cuneate: lamina chartaceous. 
glabrous: midrib sunken above: lateral veins prominulous to prominent 
below, 8-10 pairs, inarching and anastomosing to form a faint looping 
intramarginal vein at one third of the distance from the margin to the 
midrib: intercostal venation faint, reticulate: petiole 5-10 mm long. Flowers 
unknown. Fruit solitary, sessile, globose, to 4.5 cm diam.. drying much 
wrinkled and shrivelled, glabrous. Fruit calyx accrescent, divided deeply 
into 4-5 coriaceous valvate lobes. 1.5 cm long, the main axis of each lobe 
curved upwards following the curvature of the fruit base, but the sides 
bent backwards towards each other. 

Distribution and habitat: Endemic to Borneo: Sarawak (S 15932 and S 27961). 
Lowland forest at Bintulu. 

Notes: The location of the intramarginal vein at some distance from the 
margin is reminiscent of Diospyros neurosepala Bakh.. which differs in 
having leafy calyx lobes. The fruiting calyx recalls that of D. sulcata 
Kosterm.. which differs in the absence of an intramarginal vein in the leaf. 

3. Diospyros crockerensis Ng. sp. nov. 

Arbores parvae ad 11 m altae: ramuli dense velutini. Folia mediocra. 
elliptica. 7-17 x 2.5-8.3 cm. apice acuminata, basi cuneata. lamina chartacea 
infra dense pubescentia. costa supra immersa sulco puberulo: nervi 
secundarii subtus prominentes 6-9 pares: nervi tertiarii subtus prominuli 
laxe scalariformes: petioli c. 0.5 cm longi dense velutini. Flores masculi 3-9 
vel plus ad extremos cymarum elongatarum 2^.5 cm dispositi. calyx in 
lobis 4 anguste oblongis imbricatis profunde divisus. corolla salveriformis 
ad 0.8 cm longa. Flores feminei c. 3-floris ad extremos cymarum elongatarum 
0.8-2.8 cm dispositi. calyx in lobis 4 imbricatis profunde divisus. Pedicellus 
in fructu 0.8-2.8 cm. Calyx non-accrescens. lobis c. 0.2 x 0.1 cm. Fructus 
plerumque solitarii. oblongo-ovoidei ad 3.5 x 1.5 cm. velutini. apice in 
acumine acuto gradatim contracto, pericarpio tenui lignoso. Typus: Dewol 
& Karim SAN 78389, Borneo. Sabah. Crocker Range (SAN holo. SING 
iso). 



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D , , E A B 

2 mm 

Figure 3. Diospyros crockerensis Ng. A. leafy twig; B. fruit; C. female flowers after shedding of 
corollas; D. male flower bud; E. male flower bud in longitudinal section. (A-C from SAN 
78389, D-E from S 25273). 



Diospyros (Ebenaceae) in Borneo and Peninsular Malaysia 



297 



Figure 3 

Small trees to 11 m tall. Twigs densely velvety. Leaves medium-sized, elliptic, 
7-17 x 2.5-8.3 cm. apex acuminate, base cuneate; lamina chartaceous, 
densely hairy below: midrib sunken above, the groove covered with short 
hairs: lateral veins prominent below. 6-9 pairs; intercostal venation 
prominulous below, laxly scalariform: petiole c. 5 mm long, densely velvety. 
Male flowers in clusters of 3-9 or more, at the ends of elongated 2^4.5 cm 
long cymes; calyx divided deeply into 4 narrowly oblong imbricate lobes: 
corolla salverform, to 8 mm long. Female flowers in clusters of about 3. at 
the ends of elongated 8-28 mm cymes: calyx divided deeply into 4 imbricate 
lobes. Fruits usually solitary, on 8-28 mm stalks, oblong-ovoid, with the 
apex gradually tapered to a sharp point, to 3.5 x 1.5 cm, velvety, with thinly 
woody wall. Fruit calyx non-accrescent, the lobes c. 2 x 1 mm. 

Distribution and habitat: Endemic to Borneo: Sabah (SAN 78389, SAN 
11589, SAN 130109) and Sarawak (S 25273). In hill forests. 

4. Diospyros keningauensis Ng. sp. nov. 

Arbores mediocres ad magnae. ad 40 m altae. Folia minuta ad mediocra. 
anguste elliptica ad ovata 3.5-13 x 1.5-4 cm. apice acuminata, basi cuneata 
ad rotundata. lamina chartacea glabra ad infra sparse appresse pubescentia. 
proprie rugosa in sicco. costa supra prominula in juventute appresse 
pubescens; nervi secundarii subtus prominentes 5-8 pares: nervi tertiarii 
subtus prominuli scalariformes; petioli 0.5-1 cm. Flores masculi 3-5 in 
cymis 0.5-1.5 cm dispositi. quae in quoque nodo 2 vel plus glomeratae 
possunt: calyx in lobis 4 valvatis divisus: corolla salveriformis 1.4 cm longa. 
Flos femineus solitarius pedunculo 0.5-1 cm: calyx lobis valvatis 4 (5) divisus. 
Pedicellus in fructu 0.5-1 cm. Calyx non-accrescens. lobi c. 0.5 x 0.5 cm. 
Fructus solitarius. globosus ad 4.5 cm diam.. in juventute dense pubescens. 
in sicco pericarpio crasso irregulariter rugosus. Typus: Leopold & Saikah 
SAN 74495. Borneo, Sabah (KEP holo. SAN. SING iso). 

Figure 4 

Medium and big trees to 40 m tall. Leaves small to medium-sized, narrowly 
elliptic to ovate. 3.5-13 x 1.5-4 cm. apex acuminate, base cuneate to 
rounded: lamina chartaceous. glabrous to sparsely appressed hairy below, 
drying characteristically wrinkled: midrib prominulous above, appressed 
hairy when young: lateral veins prominent below, 5-8 pairs: intercostal 
veins prominulous below, scalariform: petiole 5-10 mm long. Male flowers 



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2 mm 5 mm 



Figure 4. Diospyros keningauensis Ng. A. leafy twig; B. female inflorescence; C. female flower; 
D. female flower in longitudinal section: E. as D but after removal of ovary to show staminodes; 
F. ovary in transverse section. G. male inflorescence; H. male flower bud; I. male flower bud in 
longitudinal section; J. fruit. ( A-F from SAN 26704, B-D from SFN 27136. G-I from SFN 
27136, J from SAN 74495). 



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299 



in clusters of 3-5 on 5—15 mm long cymes, which may be clustered in 2 or 
more to each node: calyx divided into 4 valvate lobes; corolla salverform, 
c. 1.4 cm long. Female flowers solitary on a 5-10 mm peduncle; calyx 
divided into 4(-5) valvate lobes; corolla 4(-5)-lobed; staminodes 8; ovary 
bearing 8 ovules. Fruits solitary, on 5-10 mm long stalks, globose, irregularly 
wrinkled on drying, thick-walled, to 4.5 cm diam., densely hairy when 
young. Fruit calyx not accrescent, the lobes about 5x5 mm. 

Distribution and habitat: Endemic to Borneo: Sabah (Carr SFN 27136: 
SAN 26704, SAN 33783. SAN 71930. SAN 74495). Lower montane and 
montane forests at 1300-1500 m in the vicinity of Mt Kinabalu (Keningau, 
Ranau). 

5. Diospyros lunduensis Ng, sp. nov. 

Arbores mediocres ad 25 m altae. Folia mediocra. elliptica ad oblongo- 
elliptica 5.6-14 x 2.8-6 cm. apice breviter acuminata vel acuta, basi rotundata 
ad subcordata. lamina coriacea. glabra, supra in sicco ad nigra vergentia. 
costa supra immersa; nervi secundarii subtus prominentes 5-7 pares; nervi 
tertiarii supra prominuli laxe reticulati; petioli 0.5-0.8 cm. Flores masculi 
ignoti. feminei solitarii pedicellis 0.2-0.4 cm longis suffulti, calyce in lobis 4 
rotundatis imbricatis pubescentibus profunde diviso. Pedicelli in fructu 0.2- 
0.4 cm longi. Calyx non-accrescens, lobis ad 0.6 x 0.6 cm patentibus ad 
reflexis. Fructus solitarius, globosus ad 2 cm diam., glaber vel sparse appresse 
pubescens. Typus: Othman S 49996. Borneo. Sarawak. Lundu, G. Pueh 
(KEP holo. SAN iso). 

Figure 5 

Medium-sized trees to 25 m tall. Leaves medium-sized, elliptic to oblong- 
elliptic. 5.6-14 x 2.8-6 cm. apex shortly acuminate or acute, base rounded 
to subcordate; lamina coriaceous, glabrous, tending to dry black above; 
midrib sunken above; lateral veins prominent below, 5-7 pairs; intercostal 
venation prominulous below, laxly reticulate; petiole 5-8 mm long. Male 
flowers unknown. Female flowers solitary, on 2-4 mm long stalks; calyx 
deeply divided into 4 rounded imbricate hairy lobes; corolla 4-lobed; 
staminodes 4; ovary with 4 ovules. Fruits solitary, on 2-4 mm long stalks, 
globose, to 2 cm diam.. glabrous or sparsely appressed hairy. Fruit calyx 
not accrescent, the lobes to 6 x 6 mm, spreading to reflexed. 

Distribution: Endemic to Borneo: Sarawak (5 49996 and S 25482). 



300 



Card. Bull. Singapore 53 (2001) 




Figure 5. Diospyros lunduensis Ng. A. leafy flowering twig; B. female flower bud; C. female 
flower bud in longitudinal section; D. as C but with ovary removed to show staminodes; E. 
ovary in transverse section; F. fruit. (A-E from S 25482, F from S 49996). 



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301 



6. Diospyros multinenis Ng, sp. nov. 

Arbores mediocres ad 20 m altae. Folia magna, oblongo-lanceolata 21-32 
x 5.8-9.2 cm. apice acuminata, basi subcordata. lamina chartacea glabra, 
costa supra immersa: nervi secundarii subtus prominentes 15-17 pares: 
nervi tertiarii subtus prominentes incerte scalariformes: petioli 0.8-1 cm. 
Flores ignoti. Cymae in fructu laxe ramosae 2.5-3.5 cm longae: pedicelli 1- 
1.5 cm. Calyx patelliformis. 4-radiatus. in lobis 4 valvatis divisus marginibus 
coreaceis reflexis c. 2 cm diam. Fructus 3-5. globosi 2-3 cm diam.. glabri. 
Typus: Bitrley et al. 2726. Borneo. Kalimantan (KEP holo). 

Figure 6 

Medium-sized trees to 20 m tall. Leaves large, oblong lanceolate. 21-32 x 
5.8-9.2 cm. apex acuminate, base subcordate: lamina chartaceous. glabrous: 
midrib sunken above: lateral veins prominent below. 15-17 pairs: intercostal 
venation prominent below, vaguely scalariform: petiole 8-10 mm long. 
Flowers unknown. Fruits 3-5. with 1-1.5 cm pedicels on laxly branched 
cymes 2.5-3.5 cm long, globose to elliptic. 2-3 cm diam.. glabrous, puckered 
on drying. Fruit calyx c. 2 cm diameter divided into 4 valvate lobes, forming 
a 4-pointed plate, with the sides of the lobes coriaceous and reflexed. 

Distribution and habitat: Endemic to Borneo: Kalimantan (Burley et al. 
2472 and 2726. G. Bentuang area of W. Kalimantan). Lowland forest. 

Notes: This species is very close to Diospyros wallichii King & Gamble, 
differing in the glabrous fruits borne on individual pedicels 1-1.5 cm long, 
which arise from peduncles 1-2 cm long (adding up to an infructescence c. 
3.5 cm long excluding the fruits). Overall, the inflorescence/infructescence 
is much longer than that of D. wallichii. 

The local Malay name tuba monyet (monkey tuba) indicates that the 
fruits are used as a kind of fish poison. 

7. Diospyros parabuxifolia Ng. sp. nov. 

Arbores mediocres, ad 15 m altae. Folia minuta. obovata 1.5-2 x 0.8-1.5 
cm. apice rotundata acuta vel breviter acuminata, basi cuneata ad attenuata. 
lamina chartacea glabra, costa supra immersa ad prominens: nervi secundarii 
et tertiarii invisibiles: petioli 0.1-0.2 cm. Flores ignoti. Pedicelli in fructu 
elongati 1.5-2 cm. Calyx non-accrescens. in lobis 4 imbricatis rotundatis 
0.2 x 0.2 cm profunde divisus. Fructus solitarius. ellipsoideus c. 2 x 1 cm. 
pericarpio angusto in sicco corrugato fere glabro. Typus: Was Paie S 33017. 



302 



Gard. Bull. Singapore 53 (2001) 




Figure 6. Diospyros multinervis Ng. A. leafy twig and B. f ruit. ( A-B from Burley et al. 2726). 



Dtospyros (Ebenaceae) in Borneo and Peninsular Malaysia 



303 



Borneo, Sarawak, Lawas, Bukit Batanga (KEP holo; SAN, SAR iso). 
Figure 7 

Medium-sized trees to 15 m tall. Leaves small, obovate, 1.5-2 x 0.8-1.5 cm, 
apex rounded, acute or shortly acuminate, base cuneate to attenuate; lamina 
chartaceous, glabrous; midrib sunken to raised above; lateral veins invisible; 
intercostal venation pattern invisible; petiole 1-2 mm long. Flowers 
unknown. Fruits solitary, on elongated 1.5-2 cm long stalks, ellipsoid, c. 2 x 
1 cm, thin-walled and puckered on drying, almost glabrous. Fruiting calyx 
not accrescent, divided deeply into 4 imbricate rounded lobes, c. 2 x 2 mm. 

Distribution and habitat: Endemic to Borneo: Sarawak, Lawas and Kapit 
(S 33017, S 60062). Lower montane forests at 1030-1500 m altitude. 

Notes: This species differs from Diospyros buxifolia (Blume) Hiern mainly 
in its elongated fruit stalk. From D. graciliflora Hiern, which also has an 
elongated fruit stalk, it differs in the fruit, which is ellipsoid (not globose). 
In addition, the fruit stalk of D. graciliflora is much more slender. 



New Varieties 
1. Diospyros curranii Merr., Phil. J. Sc. Bot. 4 (1909) 306. 
var. curranii 

Synonym: D. curraniopsis Bakh., Gard. Bull. S. S. 7 (1933) 168, syn. nov.; 
types: Endert4883, Borneo, Kalimantan (BO). Elmer 21464, Borneo, Sabah 
(BO). 

var. kalimantanensis Ng, var. nov. 

A varietate typica in calyce frutescenti patenti nec reflexo multo amplificato 
quo fundibulum fructu centrali formanti differt. Typus: Veldkamp 8148, 
Borneo, Kalimantan, Tumbang Tubus S of Bukit Raya (KEP holo, BO, L 
iso). 

Distribution: Borneo, Kalimantan, at Bukit Raya (Mogea 3569, 3585, 3621; 
Veldkamp 8148). 

Notes: This differs from the typical variety in the fruiting calyx being 
outstretched instead of reflexed, and much enlarged, so that the calyx 



304 



Card. Bull. Singapore 53 (2001) 




Figure 7. Diospyros parabuxifolia. A. leafy twig and B. fruit. ( A-B from S 3301 ~ ) 



Diospyros (Ebenaceae) in Borneo and Peninsular Malaysia 



305 



forms a 5-lobed funnel with the fruit in the centre. This variety appears to 
replace var. curranii in the vicinity of Bukit Raya. so male and sterile 
collections from Bukit Raya can be assumed to belong to this variety 
rather than the type variety. 

2. Diospyros ferruginescens Bakh.. Bull. Jard. Bot. Btz. Ser. iii. 15 (1941) 
409. 

var. rufotomentosa Ng. var. nov. 

A varietate typica praecipue in foliis infra dense rufe tomentosis differt: 
folia etiam latiora (ad 5.5 cm lata) possunt, nervis secundariis 
prominentioribus distincte incurvatis. Typus: Maikin et al. SAN 132942, 
Borneo. Sabah. Lumaku FR (SAN holotype). 

Medium-sized trees to 20 m tall. Twigs densely rusty hairy. Leaves medium- 
sized, oblong-elliptic. 6.5-11.5 x 3-5.5 cm. apex acuminate, base acute: 
lamina chartaceous. densely rusty hairy below, tending to dry black above; 
midrib sunken above, the groove often filled with hairs; lateral veins 
prominent below. 6-10 pairs, inarching strongly at the margins: intercostal 
veins prominent below, reticulate: petiole c. 5 mm long. Male flowers in 
clusters of 3-9. shortly pedicelled. sharing a common 5-7 mm peduncle: 
calyx divided into 4 small triangular valvate teeth: corolla salverform. 5-7 
mm long. Female flowers not known. Fruits solitary, on 1-2 mm long stalks, 
ovoid to globose, to 1.5 cm diam.. densely rusty hairy. Fruit calyx accrescent, 
divided deeply into 4-5 triangular valvate lobes, forming a 4-5 pointed 
star, each lobe c. 5x4 mm. 

Distribution and habitat: Endemic to Borneo: Sabah (SAN 55535, SAN 66013. 
SAN 102122. SAN 113977, SAN 132942, SAN 139235), Kalimantan 
(Kostermans 8778). Lowland and hill forests up to 700 m. 

Notes: This differs from the typical variety mainly in the underside of the 
leaves, which is densely rusty-hairy. The leaves may also be broader (to 5.5 
cm wide) with the lateral veins more prominent and distinctly inarching. 

3. Diospyros lanceifolia Roxb., Hort. Beng. (1814) 93. 
var. iliaspaiei Ng. var. nov. 

Arbores mediocres ad 27 m altae; ramuli dense ferrugineo-pubescentes. 



306 



Card. Bull. Singapore 53 (2001) 



Folia mediocra ad magna, oblonga ad oblongo-lanceolata 13.5-35 x 3.5-14 
cm, apice acuminata ad acuta, basi cuneata ad rotundata interdum fovearum 
glandulosarum pare uno instructa, lamina coriacea infra dense ferrugineo- 
pubescentia inter venas bullata, costa supra immersa; nervi secundarii subtus 
prominentes, supra immersi, 15-24 paribus incurvatis ad marginem in nervo 
intramarginali conspicuo anastomosantibus; nervi tertiarii subtus 
prominentes supra immersi transverse reticulati; petioli 1-2.4 cm. Flores 
feminei c. 3, in cymis brevibus 0.2-0.5 cm dispositi. Pedunculi in fructu 
breves, 0.2-0.5 cm longi. Calyx lobis c. 1 x 0.8 cm marginibus reflexis. 
Fructus 1-3, globosi nitide nigri, ad 2 cm diam, in juventute velutini. A var. 
consanguinea in nervis secundariis laminae quoque latere in nervo 
intramarginali valde evoluto anastomosantibus differt. Typus: Mas Paie S 
50180, Borneo, Sarawak (KEP holo, SAN, SAR iso). 

Medium-sized trees to 27 m tall. Twigs densely rusty hairy. Leaves medium- 
sized to large, oblong to oblong-lanceolate, 13.5-35 x 3.5-14 cm, apex 
acuminate to acute, base cuneate to rounded, sometimes bearing a pair of 
pit-glands; lamina coriaceous, densely rusty hairy below, bullate between 
the veins; midrib sunken above; lateral veins sunken above, prominent 
below, 15-24 pairs, inarching and joining at the margins to form a prominent 
intramarginal vein; intercostal venation sunken above, prominent below, 
reticulo-transverse; petiole 1-2.4 cm long. Female flowers in clusters of c. 3 
on short 2-5 mm long cymes. Fruits 1-3, on short 2-5 mm long stalks, 
globose, shiny black, to 2 cm diam., velvety when young. Fruit calyx: lobes 
c. 10 x 8 mm, with sides reflexed. 

Distribution and habitat: Endemic to Borneo: Sarawak (S 24803, S 39109, S 
41224, S 50180, AZ 5671). Lowland forests. 

var. renageorgei Ng, var. nov. 

Arbores mediocres ad 21 m altae. Folia mediocra, elliptica ad ovata 6-19.5 
x 4-8.5 cm, apice acuminata, basi rotundata paullo attenuata saepe fovearum 
glandulosarum pare uno provisa, lamina chartacea glabra supra plerumque 
subtus interdum nitentia, inter venas paullo bullatum, costa supra immersa; 
nervi secundarii supra immersi, subtus prominentes 6-12 pares; nervi tertiarii 
supra immersi subtus prominuli transverse reticulati; petioli 0.8-1.5 cm. 
Flores feminei 1-5 in cymis ad 0.6 cm longis dispositi. Pedicelli in fructu 
0.3-0.6 cm. Calyx lobis ad 1 x 1 saltern marginibus reflexis. Fructus circa 3, 
globosi 1.3 cm diam., velutini. A var. lancei [folia in laminae basi rotundata 
nec cuneata differt. Typus: Rena George S 43068, Borneo, Sarawak, 
Limbang (KEP holo, SAN iso). 



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307 



Medium-sized trees to 21 m tall. Leaves medium-sized, elliptic to ovate. 6- 
19.5 x -4-8.5 cm. apex acuminate, base rounded, slightly attenuate, and 
often bearing a pair of pitted glands: lamina chartaceous. glabrous, usually 
shiny on the upper surface and sometimes on the lower surface, slightly 
bullate between the veins: midrib sunken above: lateral veins sunken above, 
prominent below. 6-12 pairs: intercostal venation sunken above, prominent 
below, reticulo-transverse: petiole 8-15 mm long. Female flowers in clusters 
of 1-5 on cymes up to 6 mm long. Fruits about 3. on 3-6 mm long stalks, 
globose, c. 1.3 cm diam.. velvety. Fruit calyx with lobes up to 1 x 1 cm. the 
lobes or at least their sides reflexed. 

Distribution and habitat: Endemic to Borneo: Sabah (5A.V 99668). Sarawak 
(5 17862. S 17897. S 42254. S 43068. S 47134) and Brunei (Coode 7050). 
Lowland forests. 

Sotes: Named in honour of Rena George. 1956 — 1994. botanist of the 
Sarawak Forestry Department, who sacrificed her life for the sake of 
advancing botanical knowledge. 

Specimen 5 36348 from Kapit at 950 m altitude is placed here with 
some doubt: its leaves are not shiny like the others. 

var. saliciformis Ng. var. nov. 

Arbores mediocres ad 20 m altae. Folia elongata. saliciformia 5.5-11.5 x 1- 
3.2 cm. apice gradatim contracta. basi gradatim contracta et attenuata 
(glandes desunt). lamina chartacea glabra, costa supra immersa: nervi 
secundarii prominuli ad fere invisibiles 5-7 pares: nervi tertiarii subtus fere 
invisibles: petioli 0.3-0.5 cm. Flores ignoti. Pedicelli in fructu 0.1-0.2 cm. 
Calyx stellae 4-radiatae similis. lobis ad 0.6 x 0.4 cm. Fructus solitarius. 
subsessilis. globosus ad 1.5 cm diam.. velutinus glabrescens ad glaber. A 
varietatibus ceteris in laminis angustis saliciformibus differt. Typus: Coode 
7826. Borneo. Brunei (KEP holo. BRUN. SAN. SING iso). 

Medium-sized trees to 20 m tall. Leaves elongated, willow-like. 5.5-11.5 
x 1-3.2 cm. apex gradually tapered, base gradually tapered and attenuate, 
without pit-glands: lamina chartaceous. glabrous: midrib sunken above: 
lateral veins prominulous to almost invisible. 5-7 pairs: intercostal 
venation practically invisible below: petiole 3-5 mm long. Flowers 
unknown. Fruits solitary, subsessile on 1-2 mm stalks, globose, velvety 
becoming glabrous, to 1.5 cm diam. Fruit calyx a 4-pointed woody star 
with the lobes up to 6 x 4 mm. 



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Distribution and habitat: Endemic to Borneo: Sarawak {Chew 1170,S 49201) 
and Brunei (Coode 7826). Lowland forest. 

Notes: The leaves of Diospyros lanceifolia var. saliciformis are superficially 
similar to those of D. eucalyptifolia Bakh. and the narrow-leafed riverine 
form of D. andamanica (Kurz) Bakh. 

3. Diospyros penibukanensis Bakh., Bull. Jard. Bot. Btz. Ser. iii, 15 (1937) 
124. 

var. scalarinervis Ng, var. nov. 

A var. penibukanensi in foliis subtus glabris nec dense pubescentibus, nervis 
tertiariis scalariformibus remotioribus, ramulorum et foliorum pilis tenuibus 
appressis nec patentibus differt. Varietates etiam geographicaliter disjunctae 
prima Sarawak secunda Sabah incola. Typus: S 56607, Borneo, Sarawak 
(KEP holo, SAN, SAR iso). 

Small trees to 10 m tall, rarely to 25 m. Twigs velvety on young parts. 
Leaves medium-sized to large, elliptic to broadly obovate or oblong-obovate, 
11-38 x 4-15 cm, apex acuminate, base subcordate, rarely only rounded; 
lamina chartaceous, glabrous except the veins; midrib sunken above; lateral 
veins prominulous below and finely appressed hairy, 5-24 pairs; intercostal 
venation prominent below, scalariform; petiole 8-5 mm long. Male flowers 
up to 10 in a condensed cyme c. 5 mm long; calyx with 4 imbricate triangular 
lobes. Female flowers up to 10 in a condensed cyme c. 5 mm long; calyx 
with 4 imbricate semicircular lobes. Fruits in clusters of 1-3, on c. 5 mm 
long stalks, globose, to 2.5 cm diam., velvety. Fruit calyx not accrescent, 
the calyx lobes rounded, coriaceous, 4-6 x 5-8 mm. 

Distribution and habitat: Endemic to Borneo: Sarawak (S 13586, S 13681, 
S 13702, S 18639, S 34482, S 34935, S 34962, S 35271, S 41850, S 49997, 
S 56607). Lowland, hill and heath forests. 

Notes: Variety scalarinervis differs from var. penibukanensis in its leaves, 
which are glabrous below (instead of densely hairy), the scalariform 
intercostal venation being spaced further apart, and the hairs on its twigs 
and leaves being fine and appressed (instead of spreading). They are also 
geographically disjunct, the former in Sarawak and the latter in Sabah. 



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309 



Reductions 

1. Diospyros andamanica (Kurz) Bakh.. Bull. Jard. Bot. Btz. Ser. iii, 15 
(1937) 74, (1941) 382. 

Synonyms: D. tahanensis Bakh.. Gard. Bull. S.S. 7 (1933) 163, syn. nov.; 
type: Ridley s.n.. Peninsular Malaysia. Pahang (SING). D. longepedunciilata 
Kosterm.. Blumea 23 (1977) 4h2. syn. nov.; type: SAN 29807, Borneo, 
Sabah (KEP. SAN. SAR. SING). 

2. Diospyros areolata King & Gamble. J. As. Soc. Beng. 74. 2 (1905) 228. 
Synonyms: D. bantamensis Koord. et Valeton ex Bakh.. Gard. Bull. S.S. 7 
(1933) 165. syn. nov.; type: Koorders 1718. Java (BO). D. malam Bakh.. 
Gard. Bull. S.S. 7 (1933) 176, syn. nov.; types: bb. 6335. bb. 8037, Borneo, 
Kalimantan (BO). 

3. Diospyros blancoi A.DC. Prod. 8 (1844) 151. 

Synonym: D. durionoides Bakh.. Gard. Bull. S.S. 7 (1933) 169. syn. nov.; 
types: bb. 7978. 10050 and 12456, Borneo. Kalimantan (BO). 

4. Diospyros buxifolia (Blume) Hiern. Trans. Cambr. Phil. Soc. 12, 2 (1873) 
218. 

Synonym: D. elegantissima Bakh.. Gard. Bull. S.S. 7 (1933) 169, syn. nov.: 
type: Ridley 16349. Peninsular Malaysia. Pahang (SING). 

5. Diospyros elliptifolia Merr.. Phil. J. Sci. 30 (1926) 247. 

Synonym: D. kinabaluensis (Bakh.) Kosterm.. Blumea 23 (1977) 460. syn. 
nov. [basionym: D. elliptifolia forma kinabaluensis Bakh.. Bull. Jard. Bot. 
Btz. Ser. iii. 15 (1935) 233]; types: DeVoogd 1486, Sumatra (BO): Haviland 
1322. Borneo. Sabah (SING! SAR). 

6. Diospyros foxworthyi Bakh.. Gard. Bull. S.S. 7 (1933) 171. 
Synonyms: D. levigata Bakh.. Gard. Bull. S.S. 7 (1933) 175. syn. nov.; 
types: bb. 8053, 11089, 11255, Borneo. West Kalimantan (all at BO). D. 
cylindrocarpa Kosterm.. Blumea 23 (1977) 456. syn. nov.; type: SAN 7119, 
Borneo. Sabah (SAN. SING). 

7. Diospyros korthalsiana Hiern. Trans. Cambr. Phil. Soc. 12. 2 (1873) 
168. 

Synonym: D. myrmecocalyx (Hiern) Bakh.. Gard. Bull. S.S. 7 (1933) 178, 
syn. nov. [basionym: Maba myrmecocalyx Hiern. Trans. Cambr. Phil. Soc. 
12. 2 (1873) 139]; type: Beccari 3568. Borneo. Sarawak (K). 



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8. Diospyros lateralis Hiern, Trans. Cambr. Phil. Soc. 12, 2 (1873) 167. 
Synonym: D. crassipes Bakh., Gard. Bull. S.S. 7 (1933) 167, syn. nov.; type: 
Beccari 2041, Borneo (K). 

9. Diospyros mindanaensis Merr., Phil. J. Sci. 4 (1909) 309. 

Synonym: D. endertii Bakh., Gard. Bull. S.S. 7 (1933) 169, syn. nov.; type: 
Endert 4017, Borneo, Kalimantan (BO). 

10. Diospyros montana Roxb., PL Corom. 1 (1795) 37. 

Synonym: D. pubicalix Bakh., Gard. Bull. S.S. 7 (1933) 182, Bull. Jard. 
Bot. Btz. 15 (1955) Plate 39, syn. nov.; type: Hamid C.F. 3827, Peninsular 
Thailand (BKF, BO, KEP. SING). 

Notes: Diospyros pubicalix has been an enigmatic species known only from 
its type specimen. It is well illustrated by Bakhuizen (1955). The shoots 
grow in flushes from resting buds, with the leaves exhibiting a characteristic 
sequence of changes. The first leaves are small and have cordate to 
subcordate bases. They are followed by larger leaves with rounded bases 
and finally leaves with cuneate bases. On some specimens, only parts of 
the sequence are evident, misleading authors not aware of this source of 
variation to create new species. 

11. Diospyros pilosanthera Blanco, Fl. Filip. Ed. 1 (1837) 304. 
var. pilosanthera 

Synonym: D. cubica Bakh., Gard. Bull. S.S. 7 (1933) 168, syn. nov.; type: 
Beccari 830, Borneo, Sarawak (K) 

var. elmeri (Merr.) Ng, Malay. Forester 40 (1977) 238. 

Synonym: D. nidus-avis Kosterm., Blumea 23 (1977) 464, syn. nov.; type: S 

32181, Borneo, Sarawak (SAN, SAR, SING). 

12. Diospyros plectosepala Hiern, Trans. Cambr. Phil. Soc. 12, 2 (1873) 
201. 

Synonyms: D. poiensis Bakh., Gard. Bull. S.S. 7 (1933) 181, syn. nov.; type: 
Clemens 20304 (BO, SAR). D. setosa Bakh., Gard. Bull. S.S. 7 (1933) 184, 
syn. nov.; type: Haviland 2324, Borneo (K). 

13. Diospyros ridleyi Bakh., Gard. Bull. S.S. 7 (1933) 354. 

Synonym: D. dajakensis Bakh., Gard. Bull. S.S. 7 (1933) 168, syn. nov.; 
type: bb. 10490, Borneo. Kalimantan (BO). 

14. Diospyros rufa King & Gamble, J. As. Soc. Beng. 74, 2 (1905) 228. 



Diospyros (Ebenaceae) in Borneo and Peninsular Malaysia 



311 



Synonym: D. swingleri Kosterm.. Blumea 23 (1977) 172. syn. nov.: type: 5 
26969.*Borneo. Sarawak (SAR. SING). 

15. Diospyros simaloerensis Bakh.. Gard. Bull. S.S. 7 (1933) 185. 
Synonym: D. paraoesi Bakh.. Gard. Bull. S.S. 7 (1933) 180. syn. nov.; type: 
bb. 10162. Borneo. Kalimantan (BO). 

16. Diospyros styraciformis King & Gamble. J. As. Soc. Bens. 74 ? 2 (1905) 
216. 

Synonyms: D. styraciformis var. sarawakana (Bakh.) Ng. Malay. Forester 
40 (1977) 241. syn. nov. [basionym: Diospyros sarawcikana Bakh.. Gard. 
Bull. S.S. 7 (1933) 184]: type: Haviland 2316. Borneo. Kuching (K, SAR. 
SING). D. clavipes Bakh.. Gard. Bull. S.S. 7 (1933) 166. syn. nov.; type: 
Teysmann 11397. Borneo. Kalimantan (BO). 

17. Diospyros subrhomboidea King &l Gamble. J. As. Soc. Beng. 74. 2 
(1905) 209. 

Synonyms: D. confnsa Bakh.. Gard. Bull. S.S. 7 (1933) 167. syn. nov.; type: 
Baker s.n.. Singapore (KEP. SING). D. jaheri Bakh.. Gard. Bull. S.S. 7 
(1933) 173. syn. nov.; type: Jaheri 128. Borneo. Kalimantan (BO). 

18. Diospyros sumatrana Miq.. PI. Jungh. (1851-55) 203. 

Synonyms: D. monticola Kosterm.. Blumea 23 (1977) 463. syn. nov.; type: 
Clemens 32541. Borneo. Sabah (BO). D. beccarii Hiern. Trans. Cambr. 
Phil. Soc. 12. 2 (1873) 204. svn. nov.; tvpes: Beccari 2492 and 2591. Borneo 
(K). 

19. Diospyros venosa Wall, ex A. DC.. Prod. Syst. Nat. 8 (1844) 233. 
var. venosa 

Synonym: D. bintuluensis Kosterm.. Blumea 23 (1977) 453. syn. nov.: type: 
S 15912. Borneo. Sarawak (SAN. SAR). 

var. olivacea (King & Gamble) Ng, Malay. Forester 40 (1977) 246. 
Synonym: D. olivacea (King & Gamble) Kosterm.. Blumea 23 (1977) 465. 
syn. and stat. nov., [basionym: Maba olivacea King & Gamble in Williams. 
Bull. Herb. Boiss. II. 5 (1905) 227]: type: King s Collector 7877. Peninsular 
Malaysia. Perak. Batang Padang (K. SING). 

20. Diospyros wallichii King & Gamble in Williams. Bull. Herb. Boiss. 2.5 
(1905) 429. 

Synonym: D. pulchrinervia Kosterm.. Blumea 23 (1977) 460. syn. nov.: 



312 



Gard. Bull. Singapore 53 (2001) 



type: Kostermans 21583, Borneo, Kalimantan (KEP, SAR. SING). 

Acknowledgements 

I am very grateful to the Director-General of Forest Research Institute 
Malaysia for providing research funds and facilities under the Tree Flora 
of Sabah and Sarawak Project, to the curators of the herbaria at BO, K, 
KEP, L, SAN. SAR and SING for the loan of specimens and/or the excellent 
help and facilities given during visits to their herbaria, to Dr. J.F. Veldkamp, 
National Herbarium of the Netherlands, Leiden Branch, for the Latin 
descriptions and to Mr Joseph Pao, Sarawak Forestry Department, for 
meticulously preparing the illustrations. 

Alphabetic List of Diospyros Species and Varieties Reduced 

D. bantamensis Koord. et Valeton ex Bakh. to D. areolata King & Gamble 
D. beccarii Hiern to D. sumatrana Miq. 

D. bintuluensis Kosterm. to D. venosa Wall, ex A. DC. var. venosa 

D. clavipes Bakh. to D. styraciformis King & Gamble 

D. confiisa Bakh. to D. subrhomboidea King & Gamble 

D. crassipes Bakh. to D. lateralis Hiern 

D. cubica Bakh. to D. pilosanthera Blanco, var. pilosanthera 

D. curraniopsis Bakh. to D. curranii Merr. var. curranii 

D. cylindrocarpa Kosterm. to D. foxworthyi Bakh. 

D. dajakensis Bakh. to D. ridleyi Bakh. 

D. durionoides Bakh. to D. blancoi A. DC. 

D. elegantissima Bakh. to D. buxifolia (Bl.) Hiern 

D. endertii Bakh. to D. mindanaensis Merr. 

D. jaheri Bakh. to D. subrhomboidea King & Gamble 

D. kinabaluensis (Bakh.) Kosterm. [Basionym: D. elliptifolia forma 

kinabaluensis Bakh.] to D. elliptifolia Merr. 

D. levigata Bakh. to D. foxworthyi Bakh. 

D. longepedunculata Kosterm. to D. andamanica (Kurz) Bakh. 

D. malam Bakh. to D. areolata King & Gamble 

D. monticola Kosterm. to D. sumatrana Miq. 

D. myrmecocalyx (Hiern) Bakh. [Basionym: Maba myrmecocalyx Hiern] 
to D. korthalsiana Hiern 

D. nidus-avis Kosterm. to D. pilosanthera Blanco var. elmeri (Merr.) Ng 
D. olivacea (King & Gamble) Kosterm. [Basionym: Maba olivacea King & 
Gamble] to D. venosa Wall, ex A. DC. var. olivacea (King & Gamble) Ng 



Diospyros ( Ebenaceae) in Borneo and Peninsular Malaysia 



313 



D. paraoesi Bakh. to D. simaloerensis Bakh. 

D. poiensis Bakh. to D. plectosepala Hiern 

D. pubicalix Bakh. to D. montana Roxb. 

D. pulchrinervia Kosterm. to D. wallichii King & Gamble 

D. setosa Bakh. to D. plectosepala Hiern 

D. styraciformis var. sarawakana (Bakh.) Ng [Basionym: D. sarawakana 

Bakh.] to D. styraciformis King & Gamble 

D. swingleri Kosterm. to D. rufa King & Gamble 

D. tahanensis Bakh. to D. andamanica (Kurz) Bakh. 



Gardens' Bulletin Singapore 53 (2001) 315-322. 



New and Noteworthy Records of Mosses from Mindoro, 
the Philippines, and Their Biogeographical Implication 

BENITO C TAN 

Department of Biological Sciences. National University of Singapore, 

Singapore 119260 

AND 

EMELINA H MANDIA 
Biology Department. De La Salle University. Taft Ave.. Manila. The Philippines 

Abstract 

A recent, small collection of mosses from Mt. Halcon. Mindoro Island, the Philippines, has 
yielded two new records for the country (Rhacocarpus alpinus (Wright) Par. and 
Dicranoloma daymannianum Bartr.) and eight new records for the island. Relevant 
ecological, morphological and biogeographical notes for the 15 species collected are provided. 

Introduction 

Floristic exploration was conducted from 1996 — 1997 around the summit 
of Mt. Halcon in Mindoro, the seventh largest island in the Philippines 
(Fig. 1 ). The actual summit stands at 2.582 m asl and has a grid coordinate 
of 13° 15'46" N and 120° 59'29" E. The exploration was carried out 
principally to classify and describe the vegetation types found on the 
northeastern summit zone (Mandia. 1998). In addition to a good number 
of new records of vascular plants for the island and the Philippines that 
will be reported separately by the second author, the exploration has yielded 
two species of mosses new to the country {Rhacocarpus alpinus and 
Dicranoloma daymannianum). and eight new records for the island. In the 
case of little known moss flora of Mt. Halcon. these additions are rather 
significant. Furthermore, the new records highlight the Mindoro-Palawan 
connection, a noteworthy feature of the island's biogeography. 

Summit Vegetation 

The summit vegetation of Mt. Halcon extends from 1,950 to 2,582 m altitude 
and consists largely of open, low growing species of Leptospermum. 



New moss records from Mindoro, the Philippines 



317 



podocarps. ericoids, sedges, lycopods, ferns, ground lichens and bryophytes, 
interrupted by patches of shrubs and mossy forest. Vascular cushion plants 
and geophytes. such as the sundews (Drosera), Centrolepis philippinensis 
Merr. (Centrolepidaceae), Trachymene saniculifolia Stapf (Umbelliferae), 
Aletris foliolosa Stapf (Melanthiaceae), Oreobulus ambiguous Kukenth. & 
Steenis (Cyperaceae), Patersonia lowii Stapf (Iridaceae), Gentiana 
luzoniensis Merr. (Gentianaceae), Schoenus melanostachys R. Br. and S. 
curvulus F. Muell. (Cyperaceae) abound. The soil is shallow, acidic and 
oligotrophic, deriving from mica schist and white quartz. 

Geological Origin and Biogeography of Mindoro 

Little is known about the geology and geological history of Mt. Halcon, 
except that the mountain mass consists of mainly metamorphic rocks of 
continental crust origin uplifted since mid-Miocene (Fernandez, 1982; JICA- 
MMAJ, 1984). Geologically, Mindoro. with Mt. Halcon, is interesting, 
being ascribed to be part of a Tertiary micro-continent, the North Palawan 
Block (Holloway, 1982). Others, such as Hamilton (1981), included only 
the southern Mindoro in the North Palawan Block, giving the northern 
portion a separate origin. The arrival of North Palawan Block from its 
pre-drift position contiguous with the south China mainland to its present 
day position in the Philippine archipelago has been postulated to be in 
mid- to late Pleiocene (Holloway, 1982; Hall, 1996, 1998). The event 
became a crucial step in the evolution of the modern Philippine biota 
because the resulting island chain provided the necessary land habitats for 
two-way migrations of plants and animals between Borneo and Luzon. 
Furthermore, the drifting of the ancient North Palawan Block across the 
South China Sea might also have carried with it some continental Asiatic 
plants and animals, which have become incorporated into the Philippine 
biota (Tan et al, 1988). Seen in this light, the study of Mindoro mosses 
becomes relevant in understanding the biogeography of the island. 

Floristic Affinity of Mindoro Mosses 

Floristically, the vascular plant flora of Mindoro has been identified more 
with the Luzon flora than with the Palawan flora (Merrill, 1928). Its 
incompletely known moss flora also supported a strong Luzon connection 
(Tan and Iwatsuki. 1991). It is therefore of phytogeographical significance 
that Rkacocarpus alpinus, Dicranoloma daymannianum and Acroporium 
johannes-winkleri were identified among the new collection of mosses from 
Mt. Halcon. These three Malesian taxa are widespread in distribution, at 



318 



Card. Bull. Singapore 53 (2001) 



least in Borneo, but had either no prior Philippine record in the case of R. 
alpinus and D. daymannianum, or were not previously known from 
Mindoro, like Acroporium johannes-winkleri Broth. For all three taxa, 
Mt. Halcon represents the northernmost penetration of their ranges in 
Malesia north of the equator. 

Rhacocarpus is a widespread austral genus of 7 species (Frahm, 1996), 
with R. alpinus distributed from Sumatra, Borneo, Sulawesi to New Guinea 
(Koponen and Norris, 1986). Its presence on Mt. Halcon shows the 
Gondwana influence on the mountain flora, albeit a tenuous one, which 
hitherto had not been apparent. The Mt. Halcon record also foretells the 
likely presence of this species in Palawan. Likewise, the new Mindoro 
record of Acroporium johannes-winkleri reinforces the important role of 
Palawan as a Pleistocene land bridge in the island hopping of mosses in the 
area. Earlier, Tan (1994) reported Acroporium johannes-winkleri from 
two localities in Palawan, and one locality each from Mindanao (Mt. Apo) 
and Sibuyan Island (Mt. Giting-Giting). 

It is worthwhile to note that eight moss taxa reported from Mindoro 
(see below) have no Palawan record. Under-collection is one reason for 
this phenomenon. Compared with Mindoro, however, Palawan has a better 
known moss flora owing to the four recent explorations undertaken between 
1987 — 1993 by the senior author (Tan, 1996). An alternative explanation 
for the dissimilar distribution of taxa in these two islands may lie in the 
relatively "short" time available for the full exchange of biota since the 
late Pliocene when the two islands drifted close to each other geographically. 

Annotation of the new collection of Mt. Halcon mosses 

Below we present the list of 15 moss species collected from Mt. Halcon, 
together with brief comments on their morphology and ecology. The 
collection numbers belong to the second author, with voucher specimens 
deposited at SINU and herbarium of De La Salle University-Manila campus. 

The single asterisk (*) indicates a new record for Mindoro, and double 
asterisk (**), new to the Philippines. 

* Acroporium johannis-winkleri Broth. [EHM 78]. Epiphytic on branches, 
this is a medium-sized Acroporium with rather stiff-looking and 
spreading leaves. Stems measure to 2 cm tall. It is a West Malesian 
endemic, being common in Peninsular Malaysia and Borneo. 

Braunfelsia dicranoides (Dozy & Molk.) Broth. [EHM 82]. This 
dicranaceous moss is easily recognized among its congeners by the 



New moss records from Mindoro, the Philippines 



319 



strongly falcate, non-plicate and unicostate leaves. Its long, sheathing 
perichaetial leaves are also diagnostic. It is a common mossy forest 
epiphyte throughout Malesia. 

*Breutelia arundinifolia (Duby) Fleisch. [EHM 76, 83]. This large moss is 
recognized by its big antheridial head, widely spreading to somewhat 
squarrose and sheathing leaves, and tomentose stems (cf. Virtanen. 
1997). Widespread in East Asia and Oceania, the species is a 
common ground dweller on open sites at high elevations in the 
Philippines. It has no Palawan record (Tan, 1996). 

*Campylopus exasperatus (Nees & Blume) Brid. [EHM 77]. This is an 
easily recognized Campylopus in the Philippine mountains. The 
leaves are often broadly lanceolate, acute to blunt, with a percurrent 
and broad costa that is poorly defined. The plants from Mt. Halcon 
are large, measuring to 12 cm tall and carpeting the soil underneath 
the Falcatifolium forest and heath vegetation. 

*C. hemitrichius (C. Muell.) Jaeg. [EHM 86, 92]. Distinctive in having 
narrowly lanceolate leaves with concolorous, excurrent costa, this 
species is a ground dweller in open sites at the summit of Mt. Halcon. 
There seem to be two ecotypes on this mountain, one with erect, 
appressed leaves and the other with somewhat secund leaves. Tan 
(1983) clarified the taxonomic confusion of this species vis-a-vis other 
related Philippine congeners. At present. Campylopus hemitrichius 
has no Palawan record. 

Dicranoloma assimile (Hampe) Par. [EHM 84]. This is a widespread 
Dicranoloma in Philippine mountains growing mainly on trunks and 
branches, sometimes on soil. 

**Dicranoloma daymannianum Bartr. [EHM 85]. Found attached to the 
base of trunks and branches of shrubs in the Podocarpus- 
Falcatifolium scrub, the present species resembles a small form of 
D. braunii without the filamentous propagules. The presence of a 
central strand in stem cross-section and the short, upper laminal 
cells further distinguish it from D. braunii. Klazenga (1999) discussed 
and illustrated well this uncommon Malesian taxon. Its presence in 
Palawan can be predicted. 

Leucobryum javense (Brid.) Mitt. [EHM 81]. This is the largest Leucobryum 
found commonly in Philippine forests. 

L. sanctum (Brid.) Hampe [EHM 80]. This is another common forest 
species in Malesia including the Philippines. The opening of 
perichetial bud of this epixylic and epipetric species are thickly 
covered with highly branched rhizoids that arise mainly from the 
abaxial side, but toward the base, of inner perichaetial leaves. The 



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Card. Bull. Singapore 53 {20(H) 



larger outer perichaetial leaves, oddly, do not form any rhizoidal 
outgrowth. Yamaguchi (1993) illustrated and labeled this structure 
as a '"tomentum" without any taxonomic comment. 

* Macrothamnium javense Fleisch. [EHM 75]. The species is found attached 
to prostrate branches. The strongly spinose and toothed leaf margin 
of this species is characteristic among its congeners. All Philippine 
records, thus far, are from Luzon island. 

* Racomitriwn lanuginosum [EHM 90]. Growing in open heath vegetation 
at the summit, the species is identified by its whitish and strongly 
erose leaf apices. Mt. Halcon is the second locality in the Philippines 
for this nearly cosmopolitan moss. Its first Philippine record came 
from Mt. Giting-Giting in Sibuyan Island in central Philippines (Tan, 
1993). 

** Rhaccocarpus alpinus (Wright) Par. [EHM 72]. This is the northernmost 
locality of this species in the old world tropics. The closest population 
to Mt. Halcon is on Mt. Kinabalu in NE Borneo. Plants are abundant 
on Mt. Halcon forming extensive carpets under cushion-forming 
seed plants. Its presence at high mountain peaks in Palawan should 
be sought for. 

Schlotheimia wallisii C. Muell. [EHM 91]. Epiphytic on trunk bases and 
prostrate branches, this Malesian endemic is known from Borneo, 
the Philippines and New Guinea (Vitt et a/., 1993). The other 
Philippine localities include Luzon, Negros and Mindanao, but not 
Palawan. 

Sphagnum junghuhnianum Dozy & Molk. [EHM 88]. Plants are abundant 
on wet soil under the thick growth of Miscanthus grasses. It is the 
most common species of Sphagnum in Philippine mountains. 

S. sericeum C. Muell. [EHM 87]. This is the other species of Sphagnum 
found on wet soil beneath heath vegetation on Mt. Halcon. In the 
Philippines, this species is known also from Negros and Mindanao. 

Acknowledgements 

We are grateful to Dr. R. Seppelt for his help in making available the 
article on a revision of Rhacocarpus by J.-P. Frahm. We are equally thankful 
to Dr. N. Klazenga for his critical review of the manuscript. 



Mew moss records from Mindoro. the Philippines 



321 



References 

Fernandez. J.C. 1982. Geology and Mineral Resources of the Philippines. 
Vol. One, Geology. Bureau of Mines and Geo-Sciences. Ministry of 
Natural Resources. Manila, the Philippines. 

Frahm. J. -P. 1996. Revision der Gattung Rhacocarpus Lindb. (Musci). 
Cryptogamie, Bryologique et Lichenologique. 17: 39 — 65. 

Hall. R. 1996. Reconstructing Cenozoic SE Asia. In: R. Hall & D. Blundell 
(eds.) Tectonic Evolution of Southeast Asia. The Geological Society. 
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Hall. R. 1998. The plate tectonics of Cenozoic SE Asia and the distribution 
of land and sea. In: R. Hall and J.D. Holloway (eds.). Biogeography 
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Hamilton. W. 1981. Tectonics of the Indonesian Region (second printing). 
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Holloway. N.H. 1982. North Palawan Block. Philippines - its relation to 
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Association of Petroleum Geologists Bulletin. 66: 1355 — 1383. 

JICA-Metal Mining Agency of Japan (MMAJ). 1984. Report on the 
Geological Survey of Mindoro Island. Philippine Bureau of Mines. 
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Klazenga. N. 1999. A revision of the Malesian species of Dicranoloma 
(Dicranaceae. Musci). Journal of Hattori Botanical Laboratory. 87: 1 — 
130. 

Koponen. T. and D. H. Norris. 1986. Bryophyte flora of the Huon Peninsula. 
Papua New Guinea. XVII. Grimmiaceae. Racopilaceae and 
Hedwigiaceae (Musci). Acta Botanical Fennica. 133: 81 — 106. 

Mandia. E.H. 1998. The Vegetation on the Northeastern Summit Zone of 
Mt. Hulcon. Mindoro Island. Philippines. PhD Dissertation. University 
of the Philippines at Los Banos. Laguna. 210 pp. 

Merrill. E.D. 1928. Flora of the Philippines. In: R.E. Dickerson. Distribution 
of Life in the Philippines. Bureau of Printing. Manila, the Philippines, 
pp. 130—167. 



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Tan, B.C. 1983. The status of Campylopus hemitrichius (C. Muell.) Jaeg. 
Cryptogamie, Bryologique et Lichenologique. 4: 357 — 361. 

Tan, B.C. 1993. Noteworthy range extension of Malesian mosses. Journal 
of Hattori Botanical Laboratory. 74: 227 — 233. 

Tan, B.C. 1994. The bryophytes of Sabah (North Borneo) with special 
reference to the BRYOTROP transect of Mount Kinabalu. XIX. The 
genus Acroporium (Semaophyllaceae, Musci) in Borneo, with notes on 
species of Java and the Philippines. Willdenowia. 24: 255 — 294. 

Tan, B.C. 1996. Biogeography of Palawan mosses. Australian Systematic 
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Tan, B.C. & Z. Iwatsuki. 1991. A new annotated Philippine moss checklist. 
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Tan, B.C., Z.-H. Li and P.-C. Lin. 1988. The Hainan-Mindoro connection, 
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History Bulletin of Siam Society. 36: 7 — 15. 

Vitt, D.H., T. Koponen and D.H. Norris. 1993. Bryophyte flora of the 
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Gardens ' Bulletin Singapore 53 (2001) 323-325. 



New Combinations in Malesian Myricaceae 

I.M. TURNER 
Singapore Botanic Gardens. 1 Cluny Road. Singapore 259569 

Abstract 

In line with recent opinion favouring the splitting of the genus Myrica L. sensu law. new 
combinations in Morella Lour. (Morella esculenta (Buch.-Ham.) I.M. Turner and M. javanica 
(Blume) I.M. Turner) are provided for the two Malesian species of Myricaceae. 

Introduction 

The genus Myrica L. as traditionally circumscribed consists of some 40 — 
50 extant species of trees and shrubs ranging from the arctic to the tropics. 
The generally accepted type of the genus, circumboreal Myrica gale L.. 
and the Californian endemic Myrica hartwegii S. Watson differ significantly 
from the rest of the species. This is manifested principally in having 
inflorescences inserted at the ends of the previous year's growth rather 
than on the old wood, and smooth fruits with a pair of adnate bracteoles 
rather than papillate fruits lacking adnate bracteoles (MacDonald 1989. 
Wilbur 1994). Chromosome numbers and essential oil chemistry also 
distinguish the groups (MacDonald 1989). This division has often been 
recognised at infrageneric rank (e.g. Kubitzki 1993) but. despite long- 
standing reluctance by botanists to split Myrica. the tide seems to have 
turned to favour the recognition of at least two genera. Unfortunately as 
Myrica is typified by Myrica gale L. the generic name has to remain with 
the numerically minor element and the many warm temperate and tropical 
species have to be placed in a different genus. A proposal to conserve 
Myrica with a conserved type (Verdcourt and Polhill 1997) was made in 
order to reduce the number of necessary name changes. This was rejected 
(Brummitt 1999. p. 367) largely on the grounds that, as the split was 
unsettling to the nomenclature of important species whichever element 
was maintained as Myrica. accepted typification and nomenclatural priority 
should be upheld. 

The next oldest generic name available for the tropical segregate is 
Morella Lour., typified by Morella rubra Lour., better known as Myrica 
rubra (Lour.) Sieb. & Zucc. a relatively important fruit tree in China and 
Japan. New combinations in Morella have already been published for species 
from North American (Wilbur 1994) and Africa (Killick, Polhill and 



324 



Curd. Bull Singapore 53 12001) 



Verdcourt 1998). Even if the proposal to conserve Myrica had been accepted 
it might not have prevented the resurrection of Morella because the 
proposed conserved type was to be Myrica cerifera L.. This is one of the 
waxy-fruited North American species that are possibly generically distinct 
from the fleshy-fruited African and Asian species. 

In Malesia, two species of Myrica have long been recognised: Myrica 
esculenta Buch.-Ham. and Myrica javanica Blume. See Backer (1951) and 
Noorsiha (1996) for extensive synonymies, description, illustrations and 
notes on ecology and uses. Both species are widespread, common and 
ecologically important, as well as possessing traditional utility. Both are 
clearly best placed in Morella rather than Myrica sensu stricto and new 
combinations effecting this change are made below: 

Morella esculenta (Buch.-Ham.) I.M. Turner, comb. nov. 

Basionym: Mvrica esculenta Buch.-Ham. in D. Don, Prodr. Fl. Nep. (1825) 

56. 

Type: Nepal, Hamilton s.n. (K). 

Distribution: India and Nepal, China, Indo-China, Malesia (excluding New 
Guinea). 

Morella javanica (Blume) I.M. Turner, comb. nov. 

Basionym: Myrica javanica Blume, Bijdr. Fl. Ned. Ind. (1825) 517. 

Type: Java, Mt. Gede, Reinwardt s.n. (L). 

Distribution: Malesia except Malay Peninsula. 

References 

Backer, C.A. 1951. Myricaceae. Flora Malesiana, series I. 4: 277 — 279. 

Brummitt, R.K. 1999. Report of the committee for Spermatophyta: 48. 
Taxon. 48: 359—371. 

Killick, D.J.B., R.M. Polhil! and B. Verdcourt. 1998. New combinations in 
African Myricaceae. Kew Bulletin. 53: 993 — 995. 

Kubitzki, K. 1993. Myricaceae. In: K. Kubitzki, J.G. Rohwer & V. Bittrich 
(eds.) The Families and Genera of Vascular Plants. Vol. II Flowering 
Plants • Dicotyledones. Magnoliid, Hamamelid and Caryophyllid Families. 
Springer Verlag, Berlin, Germany. Pp. 453 — 457. 



New combinations in Malesian Myricaceae 



325 



MacDonald. A.D. 1989. The morphology and relationships of the 
Myricaceae. In: P.R. Crane & S. Blackmore (eds.) Evolution, Systematics, 
and Fossil History of the Hamamelidae, Volume 2: 'Higher Hamamelidae'. 
Clarendon Press. Oxford. U.K. Pp. 147—165. 

Noorsiha. A. 1996. Myricaceae. Tree Flora of Sabah and Sarawak. 2: 245 — 
249. 

Verdcourt. B.. and R. Polhill. 1997. (1291-1292) Proposals to conserve the 
names Myrica and Gale (Myricaceae) with conserved types. Faxon. 46: 
347—348. 

Wilbur. R.L. 1994. The Myricaceae of the United States and Canada: genera, 
subgenera, and series. Sida. 16: 93 — 107. 



Gardens' Bulletin Singapore 53 (2001) 327-341. 



Sectional Placement of Three Bornean Species of Musa 
(Musaceae) based on Amplified Fragment Length 
Polymorphism (AFLP) 

CAROL WONG 1 , RUTH KIEW 2 , ANTHONY LAMB 3 , OHN SET 2 , 
SING KONG LEE 1 , LEONG HUAT GAN 1 AND YIK YUEN GAN 1 * 

1 Natural Sciences Academic Group, National Institute of Education, Nanyang 
Technological University, 1 Nanyang Walk, Singapore 637616 

2 Singapore Botanic Gardens, 1 Cluny Road, Singapore 259569 

3 Agricultural Park. Tenom, Sabah, Malaysia 

Abstract 

The traditional approach to the classification of Musa species (Musaceae) is the separation 
into four sections (Musa, Rhodochlamys, Callimusa and Australimusa) based on chromosome 
number and morphological characters. The sectional placing of Musa beccarii N.W. 
Simmonds is still unresolved due to its unique chromosome number. The sections of two 
new species from Sabah, Malaysia, M. monticola [Hotta ex] Argent and M. suratii Argent, 
were also undetermined. This study employs Amplified Fragment Length Polymorphism 
(AFLP) as a molecular tool to determine the sectional placement of these three species 
within Musa. Eight primer combinations generated 17 genetic markers, which confirmed 
M. monticola and M. suratii as distinct species. AFLP data support the inclusion of M. beccarii 
and M. monticola in sect. Australimusa. while results show M. suratii falling between 
sect. Callimusa and sect. Australimusa suggesting that the two sections can no longer be 
maintained as distinct. 



Introduction 

Banana, Musa L. (Musaceae), is an important perennial crop cultivated in 
the tropics for food, fibre and as ornamental plants. Plagued by a complex 
genetic system of sterility, interspecific hybrid constitutions, heterozygosity 
and polyploidy (Novak, 1992), taxonomic classification of Musa is still in 
need of improvement. While the genetic diversity and classification of 
cultivated edible Musa are extensively studied (Simmonds and Shepherd, 
1955; Bhat et al, 1992; Kaemmer et al, 1992), those of wild Musa are less 
emphasised. 

The traditional approach to wild Musa classification is the separation 
of Musa into four sections (Musa Baker, Rhodochlamys Sagot, Callimusa 



For correspondence: Associate Professor Gan Yik Yuen 



328 



Card. Bull. Singapore 53 (2001) 



Cheesman and Australimusa Cheesman) based on chromosome number 
and morphological characters (Cheesman, 1947; Simmonds, 1962). A fifth 
section, sect. Ingentimusa Argent, was recognised by Argent (1976) to 
include Musa ingens N.W. Simmonds from New Guinea. Section Musa is 
the largest and most diversified section ranging from South India to Japan 
and Samoa. Most edible banana cultivars, including their progenitors 
M. acuminata Colla and M. balbisiana Colla, belong to this section. Derived 
from wild species in sect. Australimusa are the Fe'i banana cultivars mainly 
distributed in New Guinea. The popular Filipino abaca originates from M. 
te.xtilis Nee, also of this section. Section Callimusa is distributed from 
Indochina to Indonesia and consists mainly of ornamental species, the best 
known being M. coccinea Andr. Section Rhodochlamys, ranging from India 
to Indochina, also consists of ornamental species, best represented by 
M. ornata Roxb.. 

However, for some species it is not clear to which section they belong. 
The sectional placing of M. beccarii N.W. Simmonds, M. monticola [Hotta 
ex] Argent and M. suratii Argent are yet to be determined. 

Simmonds (1960) described M. beccarii from Sabah, Malaysia. He 
noted it superficially resembled M. coccinea (sect. Callimusa) but is distinct 
from this species in the bracts, male buds, fruits and seeds, which are more 
similar to species from sect. Rhodochlamys. Its chromosome number, n = 9 
(Shepherd, 1959), is unlike any other Musa species: n = 10 in sect. Callimusa 
and sect. Australimusa; n = 11 in sect. Musa and sect. Rhodochlamys; and 
n = 7 in sect. Ingentimusa, although the chromosome number n = 10 has 
also been reported for M. beccarii (Shepherd, 1959). Hence, the formal 
position of M. beccarii within Musa is still undetermined. 

Musa monticola, found in montane regions of Sabah (Mt. Kinabalu 
and the Crocker Range) is a new species of undetermined section (Argent, 
2000). Argent eliminated the possibility of it belonging to sect. Callimusa 
and sect. Rhodochlamys on the grounds of it being distinct in seed structure 
and anatomy, and the male axis habit, respectively. Instead, the polished 
bracts and absence of wax suggested placement in sect. Australimusa (Hotta, 
1987). Recent cytological examination of the species shows its chromosome 
number to be n = 10 (Jong and Argent, 2001), which supports its placement 
in sect. Australimusa. 

Another new species described by Argent (2000) is M. suratii, from 
Sabah and Sarawak, Malaysia. This species is morphologically distinct within 
Musa in possessing minute seeds, giving no clue as to its formal placement 
within Musa. Jong and Argent (2001) determined its chromosome number 
as n = 10. 

Amplified Fragment Length Polymorphism (AFLP) is a reliable and 
robust fingerprinting technique widely used in genetic diversity studies of 



Sectional placement of Bornean Miisa species 



329 



plants and animals ( Vos et al. 1995). Applications of AFLP include inferring 
phylogenetic relationships (Aggarwal et al.. 1999). analysing genetic 
diversities of populations and cultivars (Paul et al., 1997; Loh et al.. 1999). 
evaluating gene flow and dispersal (Travis et al. 1996). introgression (Tohme 
et al. 1996) and hybridisation (Beismann et al.. 1997). AFLP has the 
advantage over other molecular techniques, such as RFLP. RAPD and 
microsatellites. in that it is highly reproducible, requiring no prior knowledge 
of genome sequence, relatively fast and easy to use. and in being able to 
generate multiple loci per assay. In 1999. Crouch et al. reported 
polymorphism in Musa generated by AFLP markers to be as high as that 
obtained using microsatellites. contrary to previous reports utilising soybean 
(Powell et al. 1996). Consequently. Crouch et al. (1999) concluded that 
AFLP is most effective and suitable for genetic diversity studies of Musa. 

Hence, this study aims to resolve the sectional placement of 
M. beccarii, M. monticola and M. suratii within Musa using AFLP. (Section 
Ingentimusa could not be included in this analysis due to lack of material). 

Materials and Methods 

Plant materials 

This study employed a total of fifteen Musa species with sample sizes 
ranging from one to five (Table 1). The materials included representatives 
from four sections of Musa of both wild and cultivated origin, and from a 
variety of introductions (material of the fifth section, sect. Ingentimusa. 
was not available). The samples were collected from wild populations, and 
plants grown in the Singapore Botanic Gardens (Singapore), the Royal 
Botanic Garden Edinburgh (UK) and the Agricultural Park at Tenom 
(Sabah. Malaysia). Voucher specimens are deposited in the herbaria at 
Singapore Botanic Gardens and Royal Botanic Garden Edinburgh. 

Leaf tissue was used for AFLP analysis. The leaves were surface 
sterilised using a procedure from Zhang et al. (1997). The leaves collected 
were swirled in 95% ethanol for 1 min. 5% bleach (NaOCl) for 5 min. and 
then re-immersed in fresh 95% ethanol for 30 sec, after which they were 
blotted dry and stored in sealed plastic bags at -80°C until needed for 
DNA extraction. 

DNA extraction 

Plant DNA was extracted using the CTAB method according to Reichardt 
and Rogers (1993). Briefly, leaf tissue was pulverised using liquid nitrogen 
prior to the addition of 4 ml of Solution I (2% w/v CTAB (Sigma). 100 
mM Tris-HCl, 20 mM EDTA. 1.4 M NaCl. pH 8.0) per gram of leaf tissue 



330 



Gard. Bull. Singapore 5. 



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Sectional placement of Bomean Sfusa species 



331 



and incubated for 60 min at 65 C C. The homogenate was then extracted 
with an equal volume of chloroform/isoamyl alcohol (24:1) and centrifuaed 
at 12.000 rpm for 5 min. The upper aqueous phase was recovered and 
incubated with 1/10 volume of Solution II (10% w/v CTAB. 0.7 M NaCl). 
prewarmed to 65°C. The aqueous phase was then extracted with 1 volume 
of chloroform/isoamyl alcohol (24:1) and recovered as before. To the 
recovered aqueous phase. 1 volume of Solution III (1% w/v CTAB. 50 
mM Tris-HCl. 10 mM EDTA. pH 8.0) was added and incubated overnight 
at 37°C. The mixture was then centrifuged for 5 min at 3.500 rpm and 
supernatant removed. The DNA pellet was then re-dissolved in Solution 
IV (10 mM Tris-HCl. 0.1 mM EDTA. 1 M NaCl. pH 8.0) at 0.5 to 1 ml per 
gram starting material, followed by ethanol precipitation of the DNA. It 
was then washed with 70% ethanol. dried and re-suspended in minimal 
volume of TE buffer at 0.1 to 0.5 ml per gram starting material. 

AFLP analysis 

AFLP analysis was carried out according to Vos et al. (1995) with minor 
modifications. Restriction digests of genomic DNA with EcoRl and Mse I 
were carried out at 37°C for 1 h. Following heat inactivation of the 
restriction endonucleases. the genomic DNA fragments were ligated to 
£coRI and Msel adapters overnight at 16°C to generate template DNA 
for amplification. PCR was performed in two consecutive reactions. The 
template DNA generated was first pre-amplified using AFLP primers each 
having one selective nucleotide. The PCR products of the pre-amplification 
reaction were then used as template after 5-fold dilution in sterile water, 
for selective amplification using two AFLP primers, each containing three 
selective nucleotides. A total of eight primer combinations were used in 
this study (Table 2). The final PCR products were run on a 6% denaturing 
polyacrylamide gel in IX TBE buffer. The EcoRI primers used were not 
radioactively labelled as in the original protocol. Instead, a modified silver 
staining method was used (Loh et al., 1999). 

Data analysis 

For the diversity analysis, bands were scored as present (1) or absent (0) to 
form a raw data matrix. A square symmetric matrix of similarity was then 
obtained using Jaccard*s Similarity Coefficient [x / (y-z)]. where x is the 
number of fragments in common between two taxa, y is the total number 
of fragments scored, z is the number of fragments absent in both taxa. 
from the raw data matrix. Genetic diversity estimates (GDEs) were then 
calculated as 1 minus Jaccard*s Similarity Coefficient and used for cluster 
analysis using the Unweighted Pair Group Method with Arithmetic mean 



332 



Card. Bull. Singapore 53 (2001) 



(UPGMA) technique of the NEIGHBOR program in PHYLIP version 
3.5c (Felsenstein, 1993). 



Table 2. Sequences of the primers and adapters used for AFLP analysis 



N '.\ m i * / A hhrp vi atinn 


F TV7 vmp 


i ype 


^pniipnrp f S *" - ^ ' ^ 

jCUUClltt ^ J J 1 


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FrnU T 


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V^ A V. VJ 1 PA VJ i \ V 1 VJ V VJ 1 i\ V- v_- 


HYY 109/ FA- 


FrnR I 


A H n ntpr 


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/A /A 1 1 VJVJ I / A V VI V /A VI IV 1 /A V 


r.vY 10V MA + 

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GYY 104/ MA- 




Adapter - 


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f.YY 10V F-A 


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PYY 107/ F-A AT 


/ . ( < / IX 1 


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HYY 108/ F-A AO 


LLC XX 1 


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n aptppptappa attpapt 

VJ /A V 1 VJ V VJ 1 /A V- V /A /A 1 IV /A V. 1 


r.YY 111/ F-Aff 

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U.C L/ IXl 


Pnmpr _i_ 

I 1 1111CI T J 


P APTPiPPTAPPA ATTPAPP 

VJ /A V_ 1 VJ V_ VJ 1 rtWrtrt J 1 V /A V_ V 


r.YY 1 19/ F Afr, 

VJ I I LIZ./ C r\L VJ 


FrnH I 
ZZ-C Clxl 


I IllIILI TJ 


P APTPPPTAPPA A TTP A PP 

VJiAV I VJ V VJ 1 /A V V /A /A 1 I V rVVVJ 


pyy 1 1 V F-Afif 

VJ I I 11 J/ C /A VI V 


r r/) D I 


Pnmpr _i_ "a 


Pi APTPiPPTAPPA ATTPAPiP 

VJ /A V 1 VJ V VJ 1 /A V V /A /A 1 1 V, /A VJ V 


GYY 114/ E-AGG 


£coRI 


Primer +3 


GACTGCGTACCAATTCAGG 


GYY 106/ MP 


Msel 


Primer +1 


GATGAGTCCTGAGTAAP 


GYY 115/ M-CAA 




Primer +3 


GATGAGTCCTGAGTAACAA 


GYY 116/ M-CAC 


Msel 


Primer +3 


GATGAGTCCTGAGTAACAC 


GYY 117/ M-CAG 


Msel 


Primer +3 


GATGAGTCCTGAGTAACAG 


GYY 118/ M-CAT 


Msel 


Primer +3 


GATGAGTCCTGAGTAACAT 


GYY 119/ M-CTA 


Msel 


Primer +3 


GATGAGTCCTGAGTAACTA 


GYY 120/ M-CTC 


Msel 


Primer +3 


GATGAGTCCTGAGTAAPTP 


GYY 121/ M-CTG 


Msel 


Primer +3 


GATGAGTCCTGAGTAACTG 


GYY 122/ M-CTT 


Msel 


Primer +3 


GATGAGTCCTGAGTAACTT 


Results 



Identification of species using AFLP markers 

AFLP fingerprinting of 15 species of Musa with eight primer combinations 
(Table 2) yielded a total of 276 unambiguous bands of size 50 — 500 base 
pairs. Of these, 262 (95%) are polymorphic across all samples, while the 
number of polymorphic loci generated per primer combination is 33. Figure 
1 illustrates an AFLP profile generated using primer pair 1 (E-AAC, M- 
CAA). 

A total of 17 genetic markers unique to each species were observed 
for nine species of Musa (Table 3), including M. monticola and M. suratii. 
No unique bands distinguished M. beccarii from the rest of the Musa species. 
M. suratii is distinct due to two unique bands, while another unique marker 



Sectional placement of Bornean Musa species 



333 



1 23456789 10 11121314 M 







Figure 1. AFLP profile obtained with primer pair 1 (E-AAC, M-CAA) illustrating banding 
patterns for some species examined in this study. Lane 1: M. acuminata ssp. truncata, Lane 2: 
M. balbisiana,Lane3: M. textilis, Lane 4: M. violascens, Lane 5: M. ornata. Lane 6: M. coccinea, 
Lane 7: M. nagensium. Lane 8: M. campestris, Lane 9: M. velutina. Lane 10: M jackeyi, Lane 
11: M beccarii. Lane 12: A/, si/raf/7. Lane 13: M. monticola. Lane 14: A/, borneensis. Lane M: 
pUC19////?aII molecular weight marker. 



334 

Table 3. 



Card. Bull. Singapore 53 (2001) 

Genetic markers observed for nine species of Musa 



Primer Pair EcoRl Msel 



§■ Total 
a « .a a, S number of 

I § • •* I i «, • .« 1 umque 

g s ,| « I I £ § J § markers 

II I I .« H J I I ~ per P rimer 

pair 



1 


AAC* 


CAA** 











10 


AAG 


CAC 






. i . . . 


1 


19 


ACA 


CAG 






- - - 2 - - - 


2 


28 


ACC 


CAT 






. . i . 


1 


37 


ACG 


CTA 


1 




- 11--- 


3 


46 


ACT 


CTC 






1 - 1 - - - 1 


3 


55 


AGC 


CTG 




1 


1 2 1 - - - 1 


6 


64 


AGG 


CTT 






. . . . i 


1 



Total 1 1 2 3 4 2 1 1 2 17 



*£coRI : ZscoRI-adapter based primer; the selective nucleotides added at the 3' end are 
indicated 

**Msel : Myel-adapter based primer; the selective nucleotides added at the 3' end are 
indicated 



was observed for M. monticola. The unique bands support the distinctiveness 
of both M. suratii and M. monticola as separate species. The number of 
unique bands observed for the remaining species of Musa ranged from one 
to four. 

Genetic relationships between species of Musa 

The cluster analysis using values of GDEs (Table 4) generated a phenogram 
(Fig. 2) depicting genetic relationships between species of Musa. Two main 
clusters were observed, each made up of the traditional sectional alliance 
of Musa-Rhodochlamys and Callimusa-Australirnusa respectively. The 
cluster of the Musa-Rhodochlamys alliance revealed both subspecies of M. 
acuminata, ssp. truncata and ssp. malaccensis, to be closely related as 
anticipated. M. velutina H. Wendl. & Drude and M. ornata of sect. 
Rhodochlamys clustered closely with M. acuminata, while M. nagensium 
Prain and M. balhisiana in sect. Musa formed the remaining members of 
this cluster. 

The cluster of the Callimusa-Australirnusa alliance showed M. 
violascens Ridl., M. borneensis Becc. and M. coccinea, which all belong to 
sect. Callimusa, separated from the other species in this cluster. The cluster 
was further divided into a sub-cluster consisting of M. textilis, M. beccarii, 



Sectional placement of Bornean Musa species 



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336 



Gard. Bull. Singapore 53 < 20(H) 



Musa velutina 
Musa ornata 

M. acuminata ssp. truncate 

M. acuminata ssp. malaccensis 

Musa nagensium 

Musa balbisiana 

Musa jackevi 

Musa textilis 

Musa beccarii 

Musa monticola 

Musa suratii 

Musa campestris 

Musa violascens 

Musa borneensis 

Musa coccinea 



0.26 0.13 



Figure 2. Genetic relationships of 15 species of Musa by UPGMA cluster analysis. Scale 
depicts genetic diversity estimates (GDEs). 



M. monticola, M. suratii and M. campestris Becc, while M. jackeyi Hill in 
sect. Australimusa was distant from this subcluster. 

Based on the phenogram, M. beccarii is closest to M. textilis of 
sect. Australimusa. GDE values between M. beccarii and both species of 
sect. Australimusa examined ranged from 0.249 (between M. beccarii and 
M. textilis) to 0.391 (between M. beccarii and M. jackeyi), as compared 
with species of sect. Callimusa where values ranged from 0.336 (between 
M. beccarii and M. campestris) to 0.446 (between M. beccarii and 
M. violascens). These data suggest M. beccarii to be closer genetically to 
species of sect. Australimusa than to those of sect. Callimusa. 

Musa monticola showed a similar pattern in that it was also genetically 
most similar to M. textilis. GDE values between M. monticola and species 



Sectional placement of Bornean Musa species 



337 



of sect. Australimusa ranged from 0.276 (between M. monticola and M. 
textilis) to 0.462 (between M. monticola and M. jackeyi). while values for 
species of sect. Callimusa ranged from 0.410 (between M. monticola and 
M. campestris) to 0.491 (between M. monticola and M. coccinea). This, 
too. suggests a higher similarity between M. monticola and members of 
sect. Australimusa than between M. monticola and members of sect. 
Callimusa. 

Musa suratii clustered closely with M. campestris, which belongs to 
sect Callimusa and had GDE values ranging from 0.357 (between M. suratii 
and M. campestris) to 0.526 (between M. suratii and M. borneensis) when 
compared with species in sect. Callimusa. On the other hand, species of 
sect. Australimusa possessed GDE values of between 0.358 (between 
M. suratii and M. textilis) to 0.405 (between M. suratii and M. jackeyi). GDEs 
between M. suratii and M. textilis. and M. suratii and M. campestris are 
highly similar, giving no indication of whether M. suratii is genetically closer 
to members of sect. Callimusa or sect. Australimusa. 

Discussion 

AFLP is highly informative and reliable in providing valuable insight into 
genetic similarities between species. In addition, it is useful in generating 
unique molecular markers for identification purposes and the development 
of these is important in the improvement of banana breeding programmes. 
In assessing unique bands, results confirm both M. monticola and M. suratii 
to be distinct species as described by Argent (2000). 

The separation of the Rhodochlamys-Musa and Callimusa- 
Australimusa taxa in this molecular result is in agreement with previous 
morphological data and supports the traditional chromosomal grouping of 
Musa species into the n = 11 group and n = 10 group respectively. 

Musa beccarii falls within a cluster comprising M. textilis and 
M. monticola. Simmonds and Weatherup (1990) suggested that M. beccarii 
belonged in sect. Callimusa based on morphological characters. Our results 
disagree with that placement as genetically its alliance with 
sect. Australimusa is much stronger. 

In addition, the chromosome numbers reported for M. beccarii are 
n = 9 and n = 10. this latter number being due to multivalent formation in 
meiosis (Shepherd. 1959) and which supports the inclusion of M. beccarii 
in sect. Australimusa. The subglobose seed of M. beccarii resembles that of 
sect. Australimusa and is unlike the cylindrical seed found in sect. Callimusa. 
Based on its seed structure, chromosome number and the AFLP results 
obtained. M. beccarii is confidently placed within sect. Australimusa. 



338 



Card. Bull. Singapore 53 (2001) 



Argent (2000) suggested the inclusion of M. monticola in 
sect. Australimusa based on morphological characters. AFLP results are in 
agreement with this placement, as M. monticola clusters closely with 
M. textilis and M. beccarii. Values of GDEs support the inclusion of 
M. monticola in sect. Australimusa, rather than in sect. Callimusa. Although 
M. monticola is distinct from M. textilis morphologically, e.g. in pseudostem 
height, the length of the male peduncle, imbrication of the male bud, and 
characteristics of the basal flower and seeds (Argent, 2000), they are 
genetically the most closely related. 

Interestingly, M. suratii clustered closely with M. campestris of sect. 
Callimusa, but GDEs indicate high genetic similarity with M. textilis 
suggesting M. suratii to be intermediate between sect. Callimusa and sect. 
Australimusa. Argent (2000) noted that its unique tiny seeds provide no 
clue as to its sectional alliance. Jong and Argent (2001) determined its 
chromosome number as n = 10, which further supports its position within 
the Callimusa-Australimusa group. AFLP results show M. suratii falling 
between sect. Callimusa and sect. Australimusa suggesting that the two 
sections can no longer be maintained as distinct. 

Conclusion 

AFLP has shown M. monticola and M. suratii to be distinct species as 
described by Argent (2000). In addition, results unambiguously show that 
these three species fall within the Callimusa-Australimusa alliance of Musa, 
thus disproving any notion of them belonging to the Musa-Rhodochlamys 
alliance. Their chromosome numbers also support this placement. 
M. beccarii and M. monticola possess high genetic similarities with M. textilis 
in sect. Australimusa, while M. suratii is intermediate between M. campestris 
in sect. Callimusa and M. textilis in sect. Australimusa. 

UPGMA cluster analysis illustrates why the problem has arisen as 
the three species in question, M. beccarii, M. monticola and M. suratii, in 
fact cluster between taxa in sect. Australimusa (with M. jackeyi being most 
distant) and taxa in sect. Callimusa (with M. coccinea, M. borneensis and 
M. violascens being most distant) and are intermediate between 
M. campestris in sect. Callimusa and M. textilis in sect. Australimusa. While 
M. beccarii is definitely closer to M. textilis and can therefore be said to fall 
within sect. Australimusa, M. monticola and M. suratii are intermediate 
and could equally belong to either section. The single character that 
separates the two sections is the seed type and, while that of M. monticola 
is more like that of sect. Australimusa, that of M. suratii is unique and 
conforms to neither section. This calls into question the validity of keeping 



Sectional placement of Bornean Musa species 



339 



sections Australimusa and Callimusa distinct. This problem is dealt with in 
a later study. 

Acknowledgements 

This research was funded by the Academic Research Fund, National 
Institute of Education, Nanyang Technological University, Singapore. RP 
12/98/GYY. We thank the Director of Singapore Botanic Gardens for 
permission to collect leaf samples, George Argent (Royal Botanic Garden 
Edinburgh, UK) for providing plant materials and useful comments on the 
manuscript and Kwiton Jong (University of Aberdeen, Scotland, UK) for 
making available his data on cytology. 

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Aggarwal, R.K.. D.S. Brar. S. Nandi. N. Huang and G.S. Khush. 1999. 
Phylogenetic relationships among Oryza species revealed by AFLP 
markers. Theoretical and Applied Genetics. 98: 1320 — 1328. 

Argent, G.C.G. 1976. The wild bananas of Papua New Guinea. Notes Royal 
Botanic Garden Edinburgh. 35: 77 — 114. 

Argent, G. 2000. Two interesting wild Musa species (Musaceae) from Sabah, 
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Beismann, H.. J.H.A. Barker. A. Karp and T. Speck. 1997. AFLP analysis 
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Bhat, K.V., S.R. Bhat and K.P.S. Chandel. 1992. Survey of isozyme 
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Cheesman, E.E. 1947. Classification of the bananas. II. The Genus Musa 
L. Kew Bulletin. 2: 106—117. 

Crouch. J.H.. H.K. Crouch. H. Constandt, A. Van Gysel, P. Breyne, M. 
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Jong, K. and G. Argent. 2001. Cytology of two new species of Musa 
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Kaemmer, D., R. Afza, K. Weising, G. Kahl and F.J. Novak. 1992. 
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Novak, F.J. 1992. Musa (Bananas and Plantains). In: F.A. Hammerschlag 
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Paul, S., F.N. Wachira, W. Powell and R. Waugh. 1997. Diversity and 
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A. Rafalski. 1996. The comparison of RFLP, RAPD, AFLP and SSR 
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225—238. 

Reichardt, M.J. and S.J. Rogers. 1993. Plant DNA isolation using CTAB. 
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Shepherd, K. 1959. Two new basic chromosome numbers in Musaceae. 
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Simmonds, N.W. 1960. Notes on banana taxonomy. Kew Bulletin. 14: 198 — 
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Simmonds, N.W. 1962. The Evolution of the Bananas. Longmans, London, 
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Simmonds. NAY. and K. Shephard. 1955. The taxonomy and origin of the 
cultivated banana. Journal Linnean Society (Botany). 55: 302 — 312. 

Simmonds. N.W. and S.T.C. Weatherup. 1990. Numerical taxonomy of the 
wild bananas {Musa). Sew Phytologist. 115: 567 — 571. 

Tohme. J.. D.O. Gonzalez. S. Beebe and M.C. Duque. 1996. AFLP analysis 
of gene pools of a wild bean core collection. Crop Science. 36: 1375 — 
1384. 

Travis. S.E.. J. Maschinski and P. Keim. 1996. An analysis of genetic 
variation in Astragalus cremnophylax var. cremnophylax. a critically 
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745. 

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Frijters. J. Pot. J. Peleman. M. Kupier and M. Zabeau. 1995. AFLP: a 
new technique for DNA fingerprinting. Nucleic Acids Research. 23: 
4407-^4414. 

Zhang. W.P, J.F. YVendel and L.G. Clark. 1997. Bamboozled again! 
Inadvertent isolation of fungal rDNA sequences from bamboos (Poaceae: 
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Gardens' Bulletin Singapore 53 (2001) 343-356. 



New Combinations, New Names and New Species of 
Madhuca (Sapotaceae) from Sabah and Sarawak, Borneo 

P.C. YII AND PAUL P.K. CHAI 

% Sarawak Forestry Department 
Kuching, Malaysia 

Abstract 

Five new species of Madhuca. M. engkikiana. M. markleeana. M. multinervia, M. ochracea. 
and M. silamensis. are described and two new combinations. M. daemonica and M. prolixa. 
and two new names. M. cheongiana and M. kuchingensis. are proposed as a consequence of 
the reduction of Ganua to Madhuca. 

Introduction 

In revising the family Sapotaceae for the Tree Flora of Sabah and Sarawak, 
we accepted Pennington's reduction (1991) of Ganua Dubard (1908) to 
Madhuca Buch.-Ham. ex J.F. Gmelin (1791). The enlarged genus Madhuca 
comprises 47 species and one variety in Sabah and Sarawak out of a total 
of 50 species found in Borneo. 

The Sabah and Sarawak account includes two new combinations (M. 
daemonica and M. prolixa), two new names (M. cheongiana and M. 
kuchingensis), and five new species (M. engkikiana. M. markleeana, M. 
multinervia, M. ochracea, and M. silamensis) here described. 

New Combinations and New Names 
1. Madhuca cheongiana Yii & P. Chai, nom. nov. 

Synonyms: Ganua sarawakensis Pierre ex Dubard, Bull. Mus. Hist. Nat. 
Paris 14 (1908) 409. Type: Beccari PB 3105. Borneo, Sarawak, Kuching 
(holotype FI. n.v.); Van den Assem, Blumea 7 (1953) 375; Ganua attenuata 
Griffioen & H.J. Lam. nom. ined., Anderson. A Checklist of the Trees of 
Sarawak (1980) 314, nom. mid. 

Notes: In transferring Ganua sarawakensis to the genus Madhuca. the species 
name "sarawakensis" cannot be used because it is already preoccupied by 
Madhuca sarawakensis (Pierre ex Dubard) H.J. Lam, Bull. Jard. Bot. 
Buitenz.. 3, 7 (1925) 180. based on Kakosmanthus sarawakensis Pierre ex 



344 



Card. Bull. Singapore 53 (2001) 



Dubard, Bull. Mus. Hist. Nat. Paris 14 (1908) 407, and typified by Beccari 
PB 423, Borneo, Sarawak, Kuching (holotype P, n.v.\ isotypes FI, L, n.v.). 

The species is endemic to Borneo (Brunei Darussalam, Kalimantan, 
Sabah, and Sarawak). In Sabah and Sarawak, it is uncommon and scattered 
in a few localities (SAN 29561, SAN 33601, SAN 36601 A, SAN 86191, 
SAN 126691; S 2162, S 2222, S 7559, S 15043, S 15437, S 29477, and S 
32947), and occurs mainly in lowland kerangas and mixed dipterocarp 
forests below 200 m altitude. 

The species is named in honour of Mr. Cheong Ek Choon, the 
Director of the Sarawak Forestry Department, for his interest and 
continuous support in the study of the diversity, conservation and 
sustainable management of forest tree resources in Sarawak. 

2. Madhuca daemonica (Van den Assem) Yii & P. Chai. comb. nov. 

Basionym: Ganua daemonica Van den Assem, Blumea 7 (1953) 394. Type: 
Egar A 0932, Borneo, Sarawak, Setapok FR (holotype KEP!). 

Notes: A species endemic to Borneo and found in a few localities in Sabah 
(SAN 17437, SAN 17448), and more commonly in Sarawak (Egar A 0932, 
S 2618, S 2722, S 2768, S 4423, S 4666, S 5866, S 7088, S 12393, S 12965, S 
14462, S 30039, and S 36452). Its habitats include peatswamp, kerangas 
and mixed dipterocarp forests at altitudes up to 700 m. 

3. Madhuca kuchingensis Yii & P. Chai. nom. nov. 

Based on Ganua beccarii Pierre ex Dubard, Bull. Mus. Hist. Nat. Paris 14 
(1908) 408. Type: Beccari PB 2241, Borneo, Sarawak (holotype FI, n.v.). 

Notes: In the genus Madhuca, there exists M. beccarii (Engl.) H.J. Lam, 
Bull. Jard. Bot. Buitenz., 3, 7 (1925) 177, based on Payena beccarii Engl., 
Bot. Jahrb. 12 (1890) 508, and typified by Beccari PB 1598, Borneo, 
Sarawak, Kuching (holotype FI, n.v.; isotypes K, P, n.v.). The new name 
Madhuca kuchingensis Yii & P. Chai is, therefore, proposed. 

The species is endemic to Borneo and confined to the central and 
western parts of Sarawak (S 3356, S 4307, S 14846, S 24344, S 24514, S 
24516, S 24545, S 24602, S 27064, S 32394, S 32472, S 37044, and S 37861), 
and found mainly in lowland kerangas and mixed dipterocarp forests at 
altitudes up to 100 m. 



New combinations, names and species of Madhuca in Borneo 



345 



4. Madhuca prolixa (Pierre ex Dubard) Yii & P. Chai, comb. nov. 

Basionym: Gamia prolixa Pierre ex Dubard. Bull. Mus. Hist. Nat. Paris 14 
(1908) 409. Type: Beccari PB 2446, Borneo. Sarawak, Kuching (holotype 
FI, m.v.; isotype L, n.v.). 

Notes: A species occurring in Peninsular Malaysia and Borneo. Scattered 
throughout Sarawak in lowland mixed dipterocarp forest at altitudes up to 
200 m (S 4099, S 18771. S 25258, and S 29249). It also occurs in Brunei 
(B RUN 2476 and B RUN 3317). 

New Species 
1. Madhuca engkikiana Yii & P. Chai. sp. nov. 

(Engkik Soepadmo. Coordinator and Chief Editor of the Tree Flora of 
Sabah & Sarawak project) 

Madhncae kuchingensi arete similis, sed in ramulis glabris folds bene 
dispositis, venatione intercostali reticulata supra non impressa, venis paucis 
e costa enascentibus venis lateralibus parallelis differt. Typus: Kodoh Tarodop 
SAN 83612. Borneo. Sabah. Sandakan. Telupid (holotypus SAR!: isotypi 
AA, K. KEP!. L. SAN!, SING!). 

Figure 1 

Tree up to 25 m tall, 42 cm diameter. Bark reddish grey: inner bark yellowish 
red. Sapwood yellow. Twigs slender, terete, glabrous. Terminal buds c. 3 
mm long, puberulous. Stipules triangular, c. 1 x 0.4 mm. Leaves spirally 
arranged, scattered and well-spaced along the twigs, coriaceous, glabrous 
on both surfaces: blade elliptic to elliptic-obovate, 9-13 x 3.5^1.8 cm, base 
cuneate. slightly decurrent and oblique, margin entire and plane, apex 
short-acuminate; midrib raised on both surfaces: lateral veins 11-15 pairs, 
ascending at an angle of c. 80° from the midrib, distinctly connected by the 
thickened intercostal veins to form intramarginal vein-loops, prominent on 
both surfaces; intercostal venation reticulate, with a few veins arising from 
the midrib and parallel to the lateral veins; petioles 2-3.5 cm long, flat to 
slightly raised on the adaxial side, thickened, black and puberulous at the 
base. Inflorescences axillary, 6-8-flowered; pedicels up to 0.3 cm long, 
yellowish appressed hairy. Flowers: calyx biseriate, consisting of two whorls 
of two sepals; sepals suborbicular. c. 3 x 2.5 mm, tufted hairy at the apex; 
corolla c. 2 mm long, tube c. 0.75 mm long, lobes 7. oblanceolate, densely 
pubescent with tufted yellowish hairs at the apex; stamens 16. in two whorls, 



346 



Card. Bull. Singapore 53 (2001) 




Figure 1. Madhuca engkikiana. A, leafy flowering twig; B, flower bud; C, outer sepal; D. inner 
sepal: E, longitudinal section of flower bud; F, petal; G, petals and stamens; H, stamens in 
different views; I. gynoecium; J, fruit. (A-I from SAN 83612. J from SAN 131963. ) 



New combinations, names and species of Madhuca in Borneo 



347 



filaments very short, anthers lanceolate, c. 1 x 0.5 mm, slightly hairy; ovary 
subcorneal, c. 2 mm long, hairy, 6-7-loculed, style stout, c. 1.5 mm long. 
Fruits (young) ellipsoid, laterally compressed, 1.5 x 0.6-1.2 cm, densely 
brownish tomentose. Seeds laterally compressed, pointed at both ends; 
testa thin, smooth: scar linear. 

Distribution: Endemic to Borneo. Known only from four collections from 
Sabah (SAN 53982. SAN 54205, SAN 131963, and the type). 

Ecology: In lowland mixed dipterocarp forest, up to 250 m altitude. 

Notes: Similar to Madhuca kuchingensis but differs by its glabrous twigs, 
leaves well-spaced along the twigs, and reticulate intercostal venation that 
is not impressed above and with a few shorter veins arising from midrib 
and parallel to lateral veins. 

2. Madhuca markleeana Yii & P. Chai, sp. nov. 

(Mark Lee Hua Seng, Deputy Director, Forestry Department, Sarawak) 

Madhucae bomeensi et M. saraxvakensi in characteribus vegetativis similis, 
sed in ramulis laminisque foliorum glabris, stipulis orbicularibus, sepalis 
exterioribus magnis suborbicularibus et recurvatis, pedicello fructifero 
longiore (7-8 cm longo) differt. Typus: Yii S 72728, Borneo, Sarawak, 
Bukit Meluku (holotypus SAR!; isotypi K, KEP!, L, SAN!). 

Figure 2 

Tree up to 15 m tall, 25 cm diameter, with very low buttresses. Bark 
chocolate brown with greyish green mottles, smooth to finely fissured; 
inner bark c. 4 mm thick, dull orange, granular. Twigs terete, rusty-brown 
velvety hairy at the tips, glabrescent. Terminal buds up to 8 mm long; bud- 
scales elliptic, 10-14 x 9-11 mm. Stipules lanceolate, c. 10 x 3 mm, velvety 
hairy, caducous. Leaves spirally arranged, scattered and well-spaced along 
the twigs, coriaceous, glabrous on both surfaces; blade lanceolate or oblong, 
25-35 x 7-8 cm, base broadly cuneate and slightly oblique, margin entire 
and plane, apex obtuse or acuminate with a sharp tip; midrib broadly 
crested above, prominent below; lateral veins 28-35 pairs, ascending at an 
angle of 75-85° from the midrib, straight at first and then curving and 
joining at their tips to form vein-loops rather far from the leaf margin, 
impressed above, prominent below; intercostal venation slender, scalariform, 
with at least one vein descending from the margin and parallel to the 
lateral veins, faint above, distinct below; petioles 1.5-2.5 cm long, flat on 



348 



Gard Bull. Singapore 53 (2001) 




Figure 2. Madhuca markleeana. A, fruiting leafy twig; B, fruit; C. longitudinal section of fruit; 
D. part of inflorescence; E, flower bud; F, petal; G, flower bud with sepals and a few petals 
removed; H, longitudinal section of flower bud; I, petals and stamens; J, stamens in different 
views; K, gynoecium. (A-C from S 72729, D-K from 5 72728.) 



New combinations, names and species of Madhuca in Borneo 



349 



the adaxial side, round on the abaxial side, velvety hairy, glabrescent. 
Inflorescences axillary. 3-5-flo\vered; pedicels subangular, 5-6 cm long, pale 
green, sparsely pubescent. Flowers: calyx biseriate. consisting of two whorls 
of two sepals; sepals suborbicular, c. 10 x 14 mm, outer pair with recurved 
margin, inner pair smaller, margin not recurved, pubescent; corolla white, 
c. 12 mm long, lobes 8, lanceolate. 7-8 mm long, apex acute, tube c. 4 mm 
long, slightly pubescent at the throat; stamens 19, in two whorls, filaments 
subulate, c. 5 mm long, anthers sagittate, yellowish, c. 5 mm long; ovary 
disciform, c. 3 x 5 mm, 8-loculed, pilose, style filiform, pale green, tapering 
towards stigma. Fruits ellipsoid, c. 7 x 4 cm. 1-seeded. tapering at both 
ends; pericarp thick and fleshy, green, densely brown tomentose; stalk 7-8 
cm long. Seeds ellipsoid, 4.5 x 1.2-2 cm, pointed at both ends; testa thin; 
scar narrowly linear, c. 43 x 3 mm. 

Vernacular name: Sarawak — nyatoh gasing (Malay). 

Distribution: Endemic to Sarawak; known only from two collections, the 
type and Yii S 72729, Borneo, Sarawak. Bukit Meluku (K, KEP!, L, MO, 
SAN!. SAR!. SING!). 

Ecology: Understorey tree on steep slopes of lowland mixed dipterocarp 
forest, up to 400 m altitude. 

Notes: Resembles Madhuca borneensis and M. sarawakensis in vegetative 
characters but differs in having glabrous twigs and leaves, orbicular stipules, 
large suborbicular and recurved outer sepals, and longer fruit stalk (7-8 
cm long). 

3. Madhuca multinervia Yii & P. Chai. sp. nov. 

(Latin, multi = many, nervis = nerves; alluding to the many, closely parallel 
lateral veins of the leaves) 

Madhucae elmeri arete similis in characteribus vegetativis, sed in ramulis 
novellis folds petiolisque glabris, venis lateralibus 28-45 paribus arete 
approximatis ascendentibus distinguenda. Typus: Dewol SAN 97008, Borneo. 
Sabah. Tongod district. Bt. Pantagaluang (holotypus SAR!; isotypi AA, 
BO!, K. KEP!, L, OX, SAN!, SING!). 

Figure 3 

Tree up to 20 m tall. 40 cm diameter. Bark pale brown with greyish mottles, 
smooth; inner bark pale reddish, with sticky white latex. Sapwood whitish. 



350 



Card. Bull. Singapore 53 (2001) 



Twigs slender, solid, 2-4 mm diameter, subangular with distinct stipular 
scars, rusty-brown velvety hairy at the tips, quickly glabrescent and 
becoming blackish. Terminal buds c. 5 mm long. Stipules broadly ovate, c. 
6x4 mm, truncate, rusty-brown velvety. Leaves spirally arranged and 
scattered along ends of twigs, papyraceous to subcoriaceous, glabrous or 
with appressed silvery hairs on the midrib above, glabrous or with remnants 
of appressed silvery hairs and subappressed ferrugineus hairs on the midrib 
below; blade oblong-elliptic, 16.5-19 x 5-12 cm, base obliquely cuneate to 
rounded, margin entire and plane, apex obtuse; midrib impressed and 
slightly crested above, strongly prominent and rounded below; lateral veins 
28^45 pairs, ascending at an angle of 65-85° from the midrib, straight or 
slightly curved, arching and joining into vein-loops at 1-2 mm from the 
leaf margin, impressed above, prominent below; intercostal venation 
scalariform, faint; petioles 2-4 cm long, thickened, narrowly grooved on 
the adaxial side, rounded on the abaxial side, rugose and black at the base, 
glabrous. Inflorescences axillary, 3-10-flowered; pedicels 1-2 cm long, 
slender, velvety hairy. Flowers: calyx biseriate, consisting of two whorls of 
two sepals; sepals free, imbricate, orbicular, c. 6 mm across, velvety hairy, 
inner pair thinner, crested and hairy; corolla 7-10 mm long, 8-lobed, lobes 
elliptic or ovate, c. 3.5 x 1.5 mm, apex acute, densely hairy at the throat; 
stamens 16-24, in two or three whorls, filaments very short, anthers sagittate, 
c. 3.5 mm long; ovary subconical, c. 1 mm across, 8-loculed, glabrous, style 
c. 8 mm long, glabrous. Fruits ellipsoid, up to 2.1 x 1.2 cm, 1-seeded; 
pericarp thin, glabrous; stalk up to 2 cm long. Seeds laterally compressed, 
ellipsoid to obovoid, 1.7 x 1.1 x 0.8 cm, obtuse at both ends; scar as long as 
seed, c. 5 mm wide. 

Distribution: Endemic to Borneo. Scattered throughout the eastern parts 
of Sabah (SAN 88312, SAN 93874, SAN 96918, SAN 97008, SAN 99692, 
SAN 99726, SAN 111764, SAN 124566, and SAN 133478). 

Ecology: Usually on hillsides and ridges in primary lowland mixed 
dipterocarp forest. 

Notes: Closely allied to Madhuca elmeri in vegetative characters but can be 
distinguished by its glabrous young twigs, leaves and petioles, and 28-45 
pairs of closely set ascending lateral veins. 

4. Madhuca ochracea Yii & P. Chai, sp. nov. 

(Latin, ochraceus = pale yellowish brown; referring to the indumentum) 



New combinations, names and species of Madhuca in Borneo 



351 




Figure 3. Madhuca mtdtinervia. A. flowering leafy twig: B. flower bud: C. longitudinal section 
of flower bud: D. outer sepal: E. inner sepal: F. petals: G. part of flower with petals and stamens: 
H. stamen: I. gynoecium. (All from SAN 97008.) 



352 



(hud. Bull. Singapore 53 (2001) 



Madhucae dubardii similis in foliis infra persistentiter pallide flavido-brunneo 
tomentosis, sed in folii margine recurvata, apice caudato, nervis intercostalibus 
a margine descendentibus venis lateralibus parallellis differt. Typus: Wright 
S 29130, Borneo, Sarawak, Niah National Park (holotypus SAR!; isotypi 
K, L, SAN!, SING!). 

Figure 4 

Tree up to 25 m tall, 70 cm diameter. Bark greyish brown, scaly; inner bark 
dull orange. Twigs angular, glabrous. Terminal buds c. 5 mm long. Stipules 
broadly ovate, c. 5 x 4 mm, glabrous, persistent. Leaves spirally arranged 
and crowded at the ends of twigs, coriaceous, glabrous above, covered 
with persistent pale yellowish brown tomentum below; blade elliptic or 
elliptic-obovate, 8.5-19 x 3.5-8.2 cm, base cuneate, margin recurved, apex 
caudate or acuminate, acumen c. 1 cm long; midrib shallowly grooved and 
crested above, rounded and prominent below; lateral veins slender, 26-34 
pairs, ascending at an angle of 80-85° from the midrib, curved, diminishing 
and becoming inconspicuous toward the leaf margin, faint and impressed 
above, distinct below; intercostal venation slender, descending from the 
margin and parallel to the lateral veins, laxly reticulate towards the leaf 
margin, indistinct above, faint below; petioles 2-4.8 cm long, narrowly 
grooved on the adaxial side, thickened and rugose at the base. Inflorescences 
axillary, 2-7-flowered; pedicels c. 1.2 cm long, glabrous, angular and enlarged 
at the apex. Flowers: calyx biseriate, consisting of two whorls of two sepals; 
sepals orbicular, c. 6 mm across, apex rounded, velvety hairy; corolla c. 8 
mm long, 10-lobed, lobes obovate, c. 4 x 3 mm, apex rounded and ciliate; 
stamens 16, in two whorls, filaments c. 2.5 mm long, anthers sagittate, c. 2.5 
mm long; ovary subconical, c. 2 mm across, 6-8-loculed, velvety hairy, style 
c. 7 mm long, subangular, glabrous. Fruits (immature) ovoid, c. 1 cm across, 
base rounded, apex flattened and topped by stout remnant of style, pale 
yellowish brown hairy. Seeds unknown. 

Vernacular name: Sarawak — nyatoh kelabu (Iban). 

Distribution: Endemic to Borneo. Known from only two collections from 
Sarawak, the type and S 23100 from the Mentagai Hills, Marudi, Miri 
Division. 

Ecology: In primary lowland mixed dipterocarp forest on clay loam soils, 
up to 100 m altitude. 

Notes: Similar to Madhuca dubardii in the persistently pale yellowish brown 
tomentose leaf undersurface but differs by its recurved leaf margin, caudate 




Figure 4. Madhuca ochracea. A. flowering leafy twig: B. flower bud: C. longitudinal section of 
flower bud: D. outer sepal: E. inner sepal: F. petal: G. petal and stamens: H. stamen: I. gynoecium: 
J. young fruit. (A-I from 5 29130. J from S 23100.) 



354 



Gard. Bull. Singapore 53 (2001) 



leaf apex and intercostal veins descending from the margin and parallel to 
the lateral veins. 

5. Madhuca silamensis Yii & P. Chai, sp. nov. 
(of Mt. Silam, Sabah) 

Ab ullis speciebus borneensibus cognitis in foliis crasse coriaceis argenteo- 
brunneo tomentosis secus ramos crassos subangulares dispersis distincta. 
Typus: Mujin SAN 37849, Borneo, Sabah, Mt. Silam (holotypus SAR!; 
isotypus SAN!). 

Figure 5 

Tree up to 9 m tall, 15 cm diameter. Bark shallowly fissured or scaly, 
greyish brown; inner bark brittle, with white latex. Twigs stout, subangular, 
glabrous. Terminal buds up to 8 mm long. Stipules triangular, c. 4 x 4 mm, 
crested, glabrous. Leaves spirally arranged, scattered and well-spaced along 
the twigs, thickly coriaceous, silvery brown tomentose on both sides; blade 
elliptic to oblong-elliptic, 16-25 x 6-8 cm, base narrowly cuneate, margin 
entire and plane, apex blunt or rounded; midrib raised on both sides, 
stronger below; lateralveins 15-19 pairs, ascending at an angle of 70-80° 
from the midrib, curved, diminishing and becoming inconspicuous near the 
leaf margin, distinctly raised on both surfaces; intercostal venation laxly 
reticulate; petioles 3^4 cm long, grooved on the adaxial side, thickened at 
the base, glabrous. Inflorescence axillary, 2-3-flowered; pedicels 1-1.5 cm 
long, angular and slightly enlarged at both ends. Flowers: calyx biseriate, 
consisting of two whorls of two sepals; sepals broadly ovate, 6x4 mm, 
rusty-brown tomentose (mature flowers not seen). Fruits unknown. 

Distribution: Endemic to Sabah; known only from the type collection from 
Mt. Silam, Lahad Datu district and SAN 51742 from Bt. Tawai, 
Kinabatangan district. 

Ecology: In forest on ultrabasic soils, up to 850 m altitude. 

Notes: The spirally arranged leaves and the calyx comprising two whorls of 
two sepals confirm that the specimens belong to Madhuca. The species is 
distinct from any known species of Madhuca from Borneo in having thickly 
coriaceous, silvery-brown tomentose leaves that are scattered along the 
subangular and stout glabrous twigs. 




Figure 5. Madhuca silamensis. A. flowering leafy twig; B. flower bud: C. longitudinal section 
of flower bud; D. gynoecium. (All from SAN 37849.) 



356 



Card. Bull. Singapore 53 1 201 1 1 1 



Acknowledgements 

The work was supported financially by the Tree Flora of Sabah and Sarawak 
project jointly executed by the Forest Research Institute Malaysia (FRIM), 
Sabah Forestry Department and Sarawak Forestry Department. We are 
deeply indebted to the Director General and Directors of these institutions 
for their support and encouragement. Grateful acknowledgements are also 
due to the Directors/Curators of the Herbarium Bogoriense (BO), and the 
herbaria of the Forest Research Institute Malaysia (KEP), Sabah Forestry 
Department (SAN), Sarawak Forestry Department (SAR), Royal Botanic 
Gardens, Kew (K), National Herbarium of Netherlands, Leiden Branch 
(L), and Singapore Botanic Gardens (SING) for the loan of specimens 
and/or hospitality accorded to us during our visit to their institutions. We 
. are extremely grateful to Dr. J.F. Veldkamp for his kind help in providing 
Latin translations for the diagnoses of the new species. Finally we would 
like to express our thanks to Mr. Joseph Pao of the Sarawak Forestry 
Department for diligently preparing the illustrations, and to Dr. E. 
Soepadmo, Dr. Francis S.P. Ng, and Dr. W. Vink for their constructive 
criticisms and comments on the manuscript. 

References 

Dubard, M.M.M. 1908. Les Sapotacees du groupe des Illipees. Revue 
Generale Botanique. 20: 201. 

Gmelin, J.F. 1791. Systema Naturae. 2, 1: 772 & 799. 

Pennington, T.D. 1991. The Genera of Sapotaceae. Royal Botanic Gardens, 
Kew, U.K. 



Corrigenda 



Gardens' Bulletin Singapore 52 (2000) 261-271. 

The Significance of Pollen Morphology in the Taxonomy 
of the Genus Durio (Bombacaceae) 

I. SALMA 

replace page 269 with the following: 

graveolens, D. kutejensis, D. oxleyanus, were reported as oblate spheroidal. 
Sharma (1970) showed D. lowianus and D. pinangianus had a similar pollen 
shape, but that pollen of D. griffithii and D. singaporensis were oblate and 
that of D. carinatus oblate spheroidal. 

Generally, in the majority of species the thickness of the pollen wall 
varied from 2 — 4 urn. However, the pollen wall of D. griffithii and D. 
excelsus was thin (less than 2 um) while that of D. testudinarum was thick 
(greater than 4 um). 

Size varied from 46 to 145 um for the polar diameter (P) and 50 to 
120 um for the equatorial diameter (E) (Table 2). According to the 
classification of the pollen size by Erdtman (1952), Durio pollen can be 
divided into three groups, i.e. medium (25 — 50 um), large (50 — 100 um) 
and very large (100 — 200 um). Most Durio species fell within the large 
group except for D. griffithii. which had medium to large pollen, and D. 
affinis, D. oxleyanus, D. pinangianus, D. testudinarum and D. zibethinus. 
which possessed large to very large pollen (Table 2). Similar results were 
obtained by Abang Mokhtar (1991), except that pollen of his sample of D. 
griffithii and D. acutifolius fell within the medium-sized group and D. 
graveolens in the very large group. This difference in size was probably 
due to whether herbarium or fresh specimens were used. Since there was 
variation in pollen size within a species and variation between species was 
not distinct, pollen size is therefore not a good character for distinguishing 
the species. 

Conclusions 

Only in D. testudinarum are pollen characters species-specific (Table 3) 
and the pollen of this species can clearly be distinguished by a combination 
of its large size, thick wall and verrucose exine. Although there are 
differences in pollen morphological characters between the species, 
intraspecific variation also occurs. A combination of pollen characters, 
such as the exine sculpture, size and shape, can, in some cases (Table 3), 
provide supplementary information, which should, however, be used 
together with other characters for the identification of Durio species. 



Gardens' Bulletin Singapore 53 (2001) 359-361. 



Book Review: Paul Kessler (Ed) 2000. Secondary Forest Trees of 
Kalimantan, Indonesia. A Manual to 300 Selected Species. 404 pp. MOFEC 
Tropenbos. PO Box 319. Balikpapan. Indonesia. ISBN 90-5113-044-9. 
Paperback. Price US$35. available from The National Herbarium 
Netherlands. University of Leiden Branch. PO Box 9514, 2300 RA. Leiden. 
The Netherlands. 

This book is a welcome addition to the slowly growing botanical literature 
on Borneo. The work now being undertaken on the Tree Flora of Sabah 
and Sarawak (the Malaysian part of Borneo) has shown that the floristic 
congruence between Indonesian Borneo (Kalimantan) and Malaysian 
Borneo (Sabah and Sarawak) is over 80%. Any book on the flora of 
Kalimantan is useful to Sabah and Sarawak and vice versa. 

About half of the book (p 23-198) is taken up by descriptions of the 
trees, in alphabetical order by family and species. Each description is 
followed by concise notes on habitat and ecology, distribution and uses. 
For each family, there is an identification key to the genera treated, and 
under each genus, there is an identification key to the species. The other 
half of the book (p 204-403) is taken up by line drawings, one page per 
species, illustrating about two-thirds of the 300 species. 

There is no overall key to families or to the whole secondary forest 
flora. Instead, the book offers a comprehensive list of spot characters under 
which genera are listed where applicable. For example, under spot character 
4: armed plants, are listed Cratoxylum, Flacourtia, Gmelina, Leea, Oxyceros 
and Ziziphus. Under spot character 7: white sap. are listed Alstonia, 
Artocarpus, Cerbera, Dyera, Ficus, Homalanthus, Parartocarpus, Prainea 
and Tabernaemontana. Altogether 58 spot characters are compiled. I find 
spot characters very useful, in fact, more useful than the keys. The keys 
work only for the selection of species recognised in this manual, whereas 
spot characters, working at the genus level, tend to cover all species of the 
listed genera. One can query the list at any point instead of proceeding 
along the pre-set path of a key. With slight changes, this spot character list 
can be used anywhere in SE Asia. 

The descriptions and illustrations are of a high professional standard. 
Floral dissections, tertiary venation and indumentum details have been 
omitted by the artists, but the form and habit of each plant are effectively 
depicted. From my own experience. I know that getting the drawings done 
for a flora can be a real hassle. This approach sacrifices the fine details, but 
saves a lot of time. Otherwise, it would have been very difficult to get such 
a high proportion of the species illustrated. 

The plant identified as Phyllanthus emblica is actually Phyllanthus 



360 



Curd. Bull. Singapore 53 1 20(H) 



pectinatus, as indicated by the position of the fruits towards the ends of the 
leafy twigs. In P. emblica, the fruits would be close to the base. P. emblica 
ranges across mainland Asia from India to South China but stops at Perlis 
in north Peninsular Malaysia. P. pectinatus replaces it in Sumatra and 
Peninsular Malaysia south of Perlis (Ng, F.S.P. 2000. Malaysian Naturalist 
53(3): 32 — 35). All Bornean plants previously identified as P. emblica are 
likely to be P. pectinatus. 

I expected a discussion of the nature and origin of secondary forests 
in Kalimantan, but the authors, P.J.A. Kessler, P.B. Pelser, C.E. Risdale 
and K. Sediyasa have carefully avoided this topic. Instead, they offer this 
cryptic introduction: "After the completion of our manual 'Trees of the 
Balikpapan - Samarinda area, East Kalimantan, Indonesia' the production 
of a manual to selected tree species of the secondary forest was considered 
.an urgent necessity. Deforestation in Kalimantan was proceeding at an 
alarming rate and had led to millions of hectares of more or less severely 
degraded forest. The recent immense forest fires (1997-1998) added to the 
loss and made a publication even more urgent." In this way, the authors 
try to convey a sense of urgency, but do not explain how deforestation and 
burning impact upon forests, primary and secondary, in Kalimantan. A 
discussion might have been politically difficult in a book sponsored by the 
Indonesian Ministry of Forestry and Estate Crops. 

I think that what is happening in Kalimantan is too important to 
ignore. When I went to South Kalimantan recently, I found that the forests 
there had been replaced by Imperata grassland. The sheer scale of 
grasslands, in the absence of a cattle industry, took me by surprise. In the 
humid tropics, it needs work to convert woody vegetation to grassland. 
Logging alone does not result in grasslands. In 1994, I was in West 
Kalimantan and was shocked to see that for hundreds of kilometres, fires 
were burning all over the countryside. These were individually small fires 
that did not obstruct road traffic, but the smoke and haze filled the skies. 
It was obvious that these fires were the work of the people of the 
countryside. Their little houses were spaced out so that each could lay 
claim to many hectares of land - far more land than the settlers could farm 
manually. To maintain their claims, the settlers had to keep their lands 
cleared of natural vegetation. Annual burning during the dry season was 
the only way to do this. Eventually, the woody vegetation would give way 
to grassland. Why do the settlers clear more land than they can cultivate? 
They were hoping that plantation companies would eventually acquire the 
land for oil palm, pulpwood or other industrial crops, and pay compensation. 
The loggers had indeed opened the way into the forests, but the settlers 
did the rest. In this scheme of things, secondary forests in Kalimantan are 



Book Review 



361 



not forests gradually recovering from logging, but a brief transient phase 
between forest and Imperata grassland. West Kalimantan will soon look 
like South Kalimantan and the rest of Kalimantan is not far behind. 

Francis S.P. Ng 

Kepong. Malaysia 



Gardens' Bulletin Singapore 53 (2001) 363. 



Obituary 

DR GUNNAR SEIDENFADEN 

To our great sadness we learned that Dr Gunnar Seidenfaden passed away 
in February 2001 at the respectable age of 92. His career as an orchid 
taxonomist has been a long and extremely fruitful one. He continued to be 
productive right up to his death, leaving several unfinished manuscripts. 

His impact on the taxonomy of the orchids in the Indo-Chinese to 
Malaysian region has been enormous. The core of his orchidological oeuvre 
is his revision of the orchids of Thailand. In order to have at least a chance 
of completing this project he developed his own, slightly unusual but 
certainly concise style: full bibliography, full citation of specimens, good 
illustrations but no descriptions. Making use of the information gathered 
for this project, he produced a checklist for the orchids of Indo-China, and, 
together with Jeffrey Wood, he revised R.E. Holttum's Orchids of Malaya. 
For the latter project he visited Singapore on several occasions to consult 
the Singapore Botanic Gardens' herbarium. 

Seidenfaden can be ranked among the greatest in the field of South 
East Asian orchidology, on par with Smith. Schlechter and Holttum. While 
all these persons were professional plant taxonomists; for Seidenfaden, 
however, orchid taxonomy was only one of his many fields of interest. One 
could say that he 'did* the orchids of Thailand as a hobby, which makes his 
achievements all the more impressive. 

Gunnar Seidenfaden has left an admirable scientific legacy. His death 
is much regretted. 

Reference 

Seidenfaden. G and J.J. Wood. 1992. The Orchids of Peninsular Malaysia 
and Singapore. Olsen & Olsen, Denmark. 



Jaap J. Vermeulen 

Singapore Botanic Gardens 
Singapore 



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